The Hepaticae of the Yamal Peninsula, West Siberian Arctic

Full text

Arctoa (1993) 2: 57-101
THE HEPATICAE OF THE YAMAL PENINSULA,
WEST SIBERIAN ARCTIC
ПЕЧЕНОЧНИКИ ПОЛУОСТРОВА ЯМАЛ, ЗАПАДНОСИБИРСКАЯ АРКТИКА
ALEXEY D. POTEMKIN1
А. Д. ПОТЕМКИН1
Abstract
Hepatic flora of Yamal Peninsula Includes 121 species, 2 subspecies, 22 varieties and
11 forms. The conspect includes informatoion about species distribution within the peninsula,
their habitat preferences, associated species, and for selected species also taxonomlc
comments. There is described Gymnocolea fascinifera Potemk. sp. nov. and new
combinations suggested for Cephalozielia dlvarlcata var. potystratosa (Schust. & Damsh.)
Potemk. comb. nov.
(Cephalozielia
byssacea (A.Roth) Warnst. var. polystratosa Schust. &
Damsh. and also for two forms in
Barbilophozia.
Резюме
Флора печеночников полуострова Ямал насчитывает 121 вид, 2 подвида, 22
разновидности и 11 форм. В приводимом конспекте подробно указано рас-
пространение видов на полуострове, описаны их местообитания, сопутствующие виды,
а для некоторых - даны также таксономические заметки. Описан новый вид,
Gymnocolea fascinifera Potemk. sp. nov. предложены новые комбинации для
Cephalozielia divaricata var. polystratosa (Schust. & Damsh.) Potemk. comb. nov.
(Cephalozielia byssacea (A.Roth) Warnst. var. polystratosa Schust. & Damsh. и для двух
форм в роде
Barbilophozia.
I. INTRODUCTION
1. HISTORICAL BACKGROUND AND MATE-
RIALS
The report represents the first attempt
to bring together in one place and reconsider
all available information, published and un-
published, about the Hepaticae of the Yamal
Peninsula. Until the last decade, our knowl-
edge concerning this peculiar Arctic region
was almost noneexistent. A few reports
(Arnell 1918; Ladyzhenskaja 1971;
Andrejeva 1981), contain mostly fragmentary
data on about 30 species, and several
geobotanical papers mention some common
hepatics.
The most important contribution to the
study of the Yamal flora has been made by
Olga V. Rebristaya, the leader of the Yamal
group of Polar Expedition of the V. L. Ko-
marov Botanical Institute, St. Petersburg. Be-
sides of study of vascular plants, she and her
colleagues collected bryophytes and lichens
everywhere they worked from 1973 to 1983,
and in 1990 and 1992. These materials form
the basis for the reports of A. L. Zhukova
and O. V. Rebristaya (1986, 1987) on the
Hepaticae of Belyy Island and the
Matyuiyaha (Matyuiyakha) River Region and
for my studies of Yamal liverworts (Gribova
& Potemkin 1988; Potemkin 1988, 1989,
1990 a, b, c, d, e, 1992 a, b, 1993 a, b,
Czernyadjeva & Potemkin 1993, etc.).
Moreover my own collections (July, Aug.
1988) and collections of L. I. Mel'tzer and
A. P. Popov (Institute of Problem of
Development of North, Tyumen) made in
1987,
1989 and 1990 were used for the
study. Totally about 15 000 specimens of
bryophytes and lichens were investigated for
hepatics.
- Russia 197376 St.-Petersburg,
Prof.
Popova str., 2, Botanical Institute of Russian Acad. Sci. (Россия 197376
Санкт-Петербург, проф. Попова, 2, Ботанический институт РАН).

58 A. D. POTEMKIN
Table 1. Localities of Yamal, where Hepaticae have been collected
№ Locality and its abbreviation (in italics) latitude longitude
1.
SE part of Belyy Island 73° 15' N 71° 30' E
2.
Middle Khabeiyaha River
72е
25' N 72° 10' E
3.
Upper Tambey River 7Г 45' N 70° 30' E
4.
Lower Tirvyyaha River 71° 35' N 71° 30' E
5.
Basin of right tributary of Kharasavey River - Silyaha River 71° 10' N 67° 05' E
6. Middle Matyuiyaha River 70° 55f N 70° 20' E
7.
Upper Tiutey River 70° 50' N 69° 45' E
8. Middle Venuieuo River 70° 40' N 71° 00' E
9. Geological station Bovanenkovo 70° 20' N 68° 20' E
10.
Interfluve of Tomboiyaha and Syoyaha Rivers (Tomboitosyo) 70° 15' N 69° 40' E
11.
Neromayaha River Basin 70° 10' N 69° 10' E
12.
Middle Syoyaha River, norther of Mantyto Lake 70° 05' N 72° 10' E
13.
15 km NW of Marre Sale Polar Station 69° 50' N 67° 10' E
14.
Saletayaha River Basin 69° 45' N 68° 40' E
15.
Lower Yuribeitoyaha River 69° 45' N 72° 25' E
16.
Middle Sebayaha River 69° 37' N 69° 27' E
17.
Lower Khakhayayaha River 69° 35' N 67° 35' E
18.
Middle Lyakkotosyo River 69° 30' N 71° 25' E
19.
Lower Khutyyaha River 69° 45' N 70° 30' E
20.
Middle Khevesyo River 68° 35' N 73° 20' E
21.
Middle
Yuribey
River 68° 25' N 72° 10* E
22.
Lower Laptayaha River (Mys Kamennyy) 68° 20' N 73° 15' E
23.
Erkutayaha River Basin, 10 km to N from Kharangyneto Lake 68° 15' N 69° 55' E
24.
Lower Eryaha River 68° 10' N 72° 50' E
25.
Upper Khadytayaha River 67° 35' N 70° 25' E
2. METHODOLOGICAL REMARKS
Taking into account the great malleabil-
ity of arctic liverworts and the importance of
oil-bodies characteristics for their determina-
tion, I tried to identify them alive when it
was possible. Only complex analysis of vari-
ability of morphological features, data on oil-
bodies and ecological conditions made it pos-
sible to form a notion about specific criteria
of many species. Absence of data on oil-bod-
ies of some taxa of Lophozia, Scapania, Ric-
cardia, Chiloscyphus, etc. keeps some prob-
lems still insoluble. For the analysis of vari-
ability of Yamal hepatics see Potemkin
(1990b).
3. DEFINITION OF YAMAL AND PHYSIO-
GRAPHIC REMARKS
Yamal Peninsula is situated in the
north-western part of the West-Siberian
Lowland and extends from 66° 50' to 73°
30'N and from 66° 45' to 73° 30'E,
occupying about 112 000 km2 of territory
(Sisko 1977).
Yamal is considered here in the narrow
sense, i.e., its southern limit is the southern
boundary of the peninsula (some authors
also classify southern territories adjacent to
the peninsula as Yamal). At the same time I
classify here under Yamal its northernmost
part, Belyy Island, which has the same gene-
sis and is separated from the peninsula only
by the shallow Strait of Malygin. Places were
collections have been made are in Table 1
and Fig. 1.
Yamal is unique in respect of its physio-
graphic peculiarities and as the youngest
Arctic region (Rebristaya 1989). The lowland
landscape, absence of rocky outcrops,
predominantly oligotrophic, usually acid, rar-

The Hepaticae
of
the Yamal Peninsula 59
66"
68"
70*
72"
7i»*
76"
Fig.
1.
Cited localities
of
hepatic collections
(see Tab. 1).
AT
-
Arctic tundra,
NT -
northern hypoarctic tundra,
ST
- southern hypoarctic tundra.
ely neutral soils together with
the
rigorous
climate
and
young
age of the
region cause
the poverty
of the
Yamal flora
and the
formation
of
rather monotonous tundra veg-
etation
on the
peninsula. According
to
Yurt-
sev
& al.
(1978),
the
tundra zone
of the Ya-
mal Peninsula
is
divided into three subzones,
the southern hypoarctic, northern hypoarctic,
and arctic tundras
(Fig. 1).
4. AKNOWLEDGEMENTS
I express
my
sincerest thanks
to
Drs.
A. L.
Zhukova
and 0. V.
Rebristaya
for
drawing
my
attention
to the
need
for
investi-
gation
of
Yamal hepatics
and to Drs.
O.
V.
Rebristaya,
S. A.
Gribova,
I. V.
Czer-
nyadjeva,
V. V.
Golubkov
and E. N. An-
dre jeva,
who
provided numerous collections
of Yamal bryophytes
and
lichens
for my use.
I
am
indebted
to Prof. R. M.
Schuster
for the
gift
of
specimens
of
Lophozia rubrigemma
and Barbilophozia hyperborea;
to the
Direc-
tors
and
Curators
of the
following Herbaria
for
the
loan
of
material
in
their care:
S, F,
H, TRH, NICH; to Prof. S. K.
Tcherepanov
for nomenclatural assistance
and to Dr.
N.
N.
Zabinkova
for the
Latin translation
of
the description
of
Gymnocolea fascinifera.
I
am also very grateful
to Mrs. O. V.
Khitun,
who organized
my
work
in
Yamal
in 1988.
Dr.
В. М.
Murray
is
cordially thanked
for
revising
the
English language
and
some
valuable comments
on the
manuscript.
II.
SYSTEMATIC ACCOUNT
The hepatic flora
of the
Yamal Peninsula
is
rather poor.
121
species with
2
subspecies,
22 varieties
and 11
forms applied
to 39
genera,
20
families
and 3
orders,
Jungermanniales, Metzgeriales
and
Marchantiales,
are
known
for the
peninsula.
The list, which
is
based
on
detailed analysis
of
a
huge number
of
collections should
be
rather complete.
On the
other hand, addition
of mostly rare
to
Yamal, temperate,
basiphillous
and
mountain species
as
well
as
taxa
of
such problematic groups
as
Lophozia,
Scapania
and
Riccardia, which need
an
investigation based
on
living plants,
is
quite
possible.
In
the
following conspect
the
sequence
of
taxa follows Konstantinova
& al.
(1992).
Synonyms
are
listed only
(1)
when
the
name
of
a
taxon used here
is
little-known
and (2)
to emphasize
the
author's opinion
on
synonymy
of
certain taxa.
For
taxa which
have been erroneously reported
for
Yamal
under other names, these names
are
noted
with reference
to
corresponding papers.
Frequency
of
taxa
is
noted
for
each locality

60
A. D.
POTEMKIN
or,
for
taxa with similar frequency
in all
localities
of a
subzone,
for
each subzone.
The
following scale
of
frequency
is
used
for
localities where
300 or
more specimens were
collected:
if a
taxon
was
found
in 1-2
specimens
- I; 3 - 6 - II; 7 -15 - III; in
more
than
15
specimens from similar habitats
- IV;
in more than
15
specimens from different
habitats
- V. For
comparatively poorly
in-
vestigated places, only
the
number
of
collected specimens
is
listed
in
Arabic
numerals. Literature references
are
given
only
for
reports
not
confirmed
by
specimens,
or when there
is no
collection
of a
taxon
except
the
published
one. For
names
of
subzones
the
following abbreviations
are
used:
AT
(Arctic tundra),
NT
(northern
hy-
poarctic tundra)
and ST
(southern hypoarctic
tundra).
Characteristic habitats
and
presence
of
sex-
ual
and
asexual reproduction devices
are
noted
for
each taxon.
ORDER METZGERIALES Chalaud
CODONIACEAE Klinggr.
FOSSOMBRONIA
Raddi
Fossombronia alaskana Steere
& H.
Inoue
ST:
Khevesyo
I,
Er'yaha
I.
On bare
wet
loamy soil, associated with
Jungermannia polaris,
J.
obovata, Riccia
sorocarpa subsp. arctica, Cephaloziella arc-
tica, Blepharostoma,
etc.
With mature cap-
sules.
Westernmost report
for
Eurasia. Previ-
ously known from Alaska (Steere
&
Inoue
1974),
West Greenland (Mogensen
&
Bras-
sard
1978) and
Chukotka (Afonina
&
Duda
1983).
Yamal plants
of
Fossombronia alaskana
differ somewhat from American ones (Steere
& Inoue
I.c.;
Schuster 1992a). Stems thinner,
(8)9-11(12) cells high; width/length leaf
ratio more variable, from
1:0.65 - 0.90 to (in
the rare
mod.
angustifolia)
1:1.0 - 1.4; me-
dian cells
and
spores larger,
30 -
45(60)
x 28 -
34(40)
mkm and 34 - 42 mkm
respectively; greater variability
of
number
of
lamellae intersecting spore margin,
(23-
26)28
-
32(34); elaters occasionally thinner,
2-spiral
(5)6 - 8.5 mkm,
3-spiral
- 8.5-9
mkm
in
diam.
The
only seta
has
been found
On cross section
it is
only
5-6
cells
in
diam.
A development
of
perianth lamellae
is
extremely variable.
In
Yamal plants lamellae
are mostly slightly developed
and oc-
casionally impossible
to
find
on the
surface
of perianths with mature capsules. When
capsules
and
perianths
are
absent this
species
may be
confused with Lophozia
grandiretis, especially
var.
parviretis Schust.
It differs however
by the
constant absence
of
gemmae, invariably deep purple rhizoids,
position
of
gametangia, thinner cell walls
and,
as a
consequence, more delicate texture.
PELLIACEAE Klinggr.
PELLIA
Raddi
Pellia neesiana (Gott.) Limpr.
ST:
Khutyyaha
III,
Khevesyo
III,
Er'yaha
II.
In bogs with flowing water, flood plain wil-
low stands,
and
once
on
bare soil
in
spotty
tundra. Male
and
female plants occur rather
often.
ALLISONIACEAE (Schust.
ex
Grolle)
Schljak.
CALYCULARIA
Mitt.
Calycularia laxa Lindb.
et H.
Arnell
AT:
Belyy
I,
Khabeiyaha
III,
Tambey
III,
Kharasavey
II;
NT:
Matyuiyaha
I,
Neromayaha
I,
Mantyto
II,
Khakhayayaha
I; ST:
Khutyyaha
I,
Khevesyo
III,
Er'yaha
II.
On thinly turfed
and
bare, steep, often
ni-
val,
slopes;
in
lichen tundras;
on
peat out-
crops;
on
spots
of
bare soil
in
different types
of tundra; occasionally
in
sedge-moss bogs.
Male
and
female plants frequent
but
peri-
anths
and
sporophytes develop rarely. West-
ernmost reports
for
Eurasia.
The
record
on
Belyy Island
is the
northernmost
on the
globe.
Sterile
and
female plants
of
Calycularia
laxa
are
sometimes confused with Pellia.
They differ from
all
species
of
Pellia
by
ven-
tral scales, which
are
2-4(5) cells wide
at
base
and to 10
cells long. Among
the
regional
species
of
Pellia only
P.
endiviifolia (Dicks.)
Dum. develops rather long ventral slime hairs
but they
are
uniseriate
and
shorter than
ventral scales
of
Calycularia. Moreover
C.
laxa
and P.
endiviifolia
are
very different
in
ecological behaviour:
the
former
is

The Hepaticae of the Yamal Peninsula 61
acidophilus, while the latter is basiphilous.
Confusing of female plants of C. laxa with
Pellia is sometimes possible, because of
female scales are very thin, hair-like and
inconspicuous, having similar with thallus
pigmentation (see also Potemkin 1990a).
PALLAVICINIACEAE Migula
MOERCKIA Gott.
Moerckia blyttii (Moerck) Brockm.
ST:
Khutyyaha I (Andrejeva 1981H!).
In moss-lichen tundra, small groups of
plants among lichens.
BLASIACEAE Klinggr.
BlASIA L.
Blasia pusilla L.
NT: Saletayaha 1, Khakhayayaha I; ST: Khytyyaha II,
Khevesyo II, Er'yaha II.
On bare wet loamy soil on banks of brook-
lets,
rivers and lakes. Reproduction mostly
by scale-like gemmae, very rarely by discoid
gemmae in "bottles".
Plants of exposed sites often develop pur-
plish pigmentation and conspicuous white
lines along the midrib and its branches which
give them a very peculiar appearance. The
white lines are caused, according to Macvicar
(1926:77), by a deposit of calcium oxalate.
At the base of robust plants there are
numerous white lines going from the
branches. Cross section of old parts of such
plants looks septate, i.e. with thick parts with
canals filled up by calcium oxalate and thin
ones without them. The purplish pigmented
phases often develop purplish scale-like
gemmae. Secondary pigmentation, however,
has not been reported before for the gemmae
of Blasia. Schuster (1988: 235) stressed that
even in purplish plants the gemmae are al-
ways bright green.
ANEURACEAE Klinggr.
ANEURA Dum.
Aneura pinguis (L.) Dum. var. pinguis
AT: Belyy I, Khabeiyaha I; NT: Matyuiyaha I, Bova-
nenkovo 2, Neromayaha I, Khakhayayaha I; ST:
Khutyyaha II, Khevesyo II, Er'yaha II.
On spots in different types of spotty moss
tundras, on loamy slopes, occasionally in
bogs.
Male and female plants frequent, but
plants with mature capsules rare.
Aneura pinguis var. denticulata Nees
ST:
Khutyyaha & Er'yaha (Andreeva 1981).
In homogenous sedge (Carex stans) bogs
with flowing water.
RiccARDiA S. Gray
Since all specimens of Riccardia have been
studied in a dead state, i.e. without any data
on their oil-bodies, it is difficult or impossi-
ble to identify taxa of this genus correctly
(Potemkin 1991). As a consequence, all
species are listed under the sign
"cf.".
Riccardia cf. chamaedrifolia (With.) Grolle
ST:
Khutyyaha I.
In sedge-Sphagnum tussock-pool bog, on
tussock among Sphagnum. Plants monoicous,
with immature calyptral perigynia.
In the Arctic this species is inseparable
from the recently described Riccardia lati-
frons subsp. arctica Schust. & Damsh. This
arctic subspecies of R. latifrons is charac-
terized by the same morphological variability
and ecological behaviour and differs from R.
chamaedrifolia only in the absence of oil-
bodies (Schuster 1987).
It is thus possible that the collections seen
are R. latifrons subsp. arctica (which has not
been reported for Eurasia before).
Riccardia cf. latifrons (Lindb.) Lindb.
NT: Tomboitosyo I, Khakhayayaha I; ST: Khutyyaha I,
Khevesyo I.
In moss tundras and sedge-Sphagnum wil-
low stand, at base of willow on mineral soil.
ORDER JUNGERMANNIALES Klinggr.
TRICHOCOLEACEAE Nakai
Subfam. TEMNOMOIDEAE Schust.
PSEUDOLEPICOLEA
Fulf.
St J. Tayl.
Pseudolepicoiea fryei (H. Perss.) Grolle &
Ando
AT: Belyy II, Tambey II; NT: Matyuiyaha II, Bova-
nenkovo /, Khakhayayaha I; ST: Er'yaha II, near
Khadytayaha (Ladyzhenskaja 1971).
On Sphagnum tussocks in bogs, associated
mostly with Blepharostoma trichophyllum var.
brevirete.
The species is usually represented by mod.
densifolia-colorata. Mod. parvifolia-laxifolia-
viridis was found only once (Belyy).

62 A. D. POTEMKIN
Subfam. BLEPHAROSTOMATOWEAE Grolle
BLEPHAROSTOMA (Dum.) Dum.
Blepharostoma trichophyllum (L.) Dum.
var. trichophyllum
NT: Neromayaha II, Khakhayayaha I.
In nival tundra (with perianths), on top of
watershed among lichens, on willow trunk.
Blepharostoma trichophyllum var. brevirete
Bryhn & Kaal. (B. trichophyllum var. tri-
chophyllum auct. non (L.) Dum. - Andrejeva
1981).
AT V; NT: Matyulyaha IV, Tiutey 6, Venuieuo 2, Bo-
vanenkovo S, Tomboitosyo V, Neromayaha V, Mantyto V,
Marre Sale 11, Yuribeitoyaha V, Sebayaha 3,
Khakhayayaha IV, Lyakkotosyo III; ST: Khutyyaha IV,
Khevesyo V, Yuribey 4, Laptayaha V, Kharangyneto 2,
Er'yaha III, Khadytayaha 2.
Most common in diverse wet tundras with
comparatively rich soils. Rarely with perianth
and mature capsules. Plants are usually
monoicous, only exceptionally dioicous (in
Lyakkotosyo male plants were found).
Plants of this variety vary considerably in
size,
density of leaves, width of cortical cells
(20-31(40) mkm), width/length ratio of
leaf cells, shape and size of apical cells of
leaf segments. On spots in spotty tundra the
phenotypes with short leaf segments com-
posed of isodiametric cells occur. Sporadically
plants with 4 - 5 - lobed leaves and 4 - lobed
underleaves were found. Individual leaf seg-
ments of such plants are branched. Despite
great malleability, var. brevirete in Yamal is
almost always distinguished from the typical
variety by its not bulging septae, separating
the cells of leaf segments.
ANTHELIACEAE Schust.
ANTHELIA
(Dum.) Dum.
Anthelia juratzkana (Limpr.) Trev. f. ju-
ratzkana
AT: Belyy IV, Khabeiyaha III, Tambey Ш, Kharasavey
III; NT: Matyulyaha III, Tiutey /, Tomboitosyo I, Nero-
mayaha IV, Mantyto I, Saletayaha 4, Yuribeitoyaha I,
Khakhayayaha III, Lyakkotosyo I; ST: Khutyyaha III,
Khevesyo V, Laptayaha IV, Er'yaha III, Khadytayaha /.
In wet spotty tundras, on nival slopes. Of-
ten with perianths and mature capsules.
Anthelia juratzkana f. elongata Joerg.
AT: Belyy I; NT: Neromayaha I.
In spotty tundras on sandy soil, occasion-
ally together with f. juratzkana.
JUNGERMANNIACEAE Reichenb.
Subfam. LOPHOZIOIDEAE Macv.
TETRALOPHOZIA (Schust.) Schljak.
Tetralophozia setiformis (Ehrh.) Schljak.
AT: Belyy I, Khabeiyaha I, Tambey HI; NT:
Matyuiyaha II, Khakhayayaha I; ST: Laptayaha I.
In diverse tundras on sandy soil.
In Belyy and Matyuiyaha the plants of
mod. parvifolia-densifolia- (subdensifolia) -vi-
ridis (subcolorata) were found. They have
(2)3 - lobed leaves with smaller median and
especially ventral lobes and larger dorsal
ones.
In leaf form these plants resemble
Chandonanthus birmensis Steph., differing
from it in subtransverse leaf insertion, cell
net, etc. They may be confused also with
Barbilophozia quadriloba, which have, how-
ever, more symmetric, less deeply divided
leaves and a rough cuticle.
BARBILOPHOZIA Loeske sensu lato
Subgen. ORTHOCAUUS (Buch) Buch
Barbilophozia quadriloba (Lindb.) Loeske f.
quadriloba
AT: Belyy I, Khabeiyaha I, Tambey II, Kharasavey I;
NT: Matyuiyaha I, Tomboitosyo I, Marre Sale /, Sale-
tayaha 2, Yuribeitoyaha III, Lyakkotosyo I; ST: Lap-
tayaha I, Er'yaha I.
In diverse tundras with comparatively rich
soils.
Barbilophozia quadriloba f. glareosa
(Joerg.) Potemk. [Bot. Zhurn.
75(12):
1744,
1990]
AT: Belyy H, Tambey I, Kharasavey II; NT: Bova-
nenkovo У, Neromayaha II, Yurbeitoyaha II,
Khakhayayaha I; ST: Khevesyo III.
In diverse tundras, probably in less favor-
able conditions than the type form.
Barbilophozia quadriloba f. cephaloziel-
loides (Schust.) Potemk. [Novosti Sist.
Nizsh. Rast. 28:150, 1992]
AT: Belyy I.
In herb-grass-willow-moss tundra, on bare
soil, individual shoots in association with
Scapania zemljae, Lophozia heterocolpos var.
harpanthoides, Barbilophozia quadriloba
f. glareosa, Odontoschisma macounii, Junger-
mannia polaris, Cephalozia pleniceps,
Cephaloziella arctica, Blepharostoma tricho-
phyllum var. brevirete, etc.

The Hepaticae of the Yamal Peninsula 63
100 дт
1 mm
Fig. 2. Barbilophozia
hyperborea
(1-12) and B. quadriloba (13). 1, 13 - Apexes of lobes; 2-4 - Underleaves; 5-12
Leaves. Scale bars: 100 mkm - for 1, 13; 1 mm - for 2-12. All from Khutyyaha (After Polemkin 1992a).
Barbilophozia hyperborea (Schust.) R.
Stotl. & B. Stotl. ex Potemk. [Novosti Sist.
Nizsh. Rast. 28: 148, 1992] (Fig. 2).
AT:
Kharsavey I; NT: Matyuiyaha I, Tiutey Л Nero-
mayaha III, Saletayaha 2, Khakhayayaha II; ST:
Khutyyaha I, Khevesyo I.
In dwarf shrub-(herb-) moss and sedge-
moss tundras, on brook banks, mostly on
weakly turfed soil. Once (Neromayaha) with
perianths. For details and differentiation see
Potemkin (1992a).
Barbilophozia kunzeana (Hueb.) K. Muell.
f. kunzeana
AT:
Belyy I, Khabeiyaha I, Tambey II, Kharasavey II;
NT:
Matyuiyaha V, Tiutey 3, Venuieuo I, Bovanenkovo
3,
Tomboitosyo V, Neromayaha III, Marre Sale 2, Sale-
tayaha /, Yuribeitoyaha II, Sebayaha 2, Khakhayayaha
II,
Lyakkotosyo II; ST: Khutyyaha V, Khevesyo V,
Yuribey 3, Laptayaha IV, Kharangyneto 5, Er'yaha IV,
Khadytayaha 3.
In diverse moss tundras, bogs, peat out-
crops,
occasionally in late snow areas. Very
rarely with perianths and mature capsules,
sometimes with gemmae. Gemmae predomi-
nantly rhombic, 10 - 17 x 18 - 23 (31) mkm,
varying in color from green and yellowish to
brown, red brown and, occasionally, deep
purple (found in plants of mod. parvifolia-
colorata, Matyuiyaha). Leaf cells of
gemmif-
erous shoots contain (6)8 - 13(15) oil-bodies
and of shoots, not developing gemmae, (3)4 -
6 oil-bodies (found in plants from Khevesyo).
Occasionally, gemmiparous leaves have a
very rough, Barbilophozia quadriloba-like

64 A. D. POTEMKIN
cuticle (as in plants from Mantyto).
The variability of the species was described
in detail by Schuster (1969). Some pheno-
types,
however, have not been found before.
Among them: 1) plants with flat, compara-
tively narrow, not folded, appressed to stem
leaves with not reflexed sinus and small
lanceolate underleaves; 2) plants similar to
"f. plicata", differing in the absence or very
weak development of amphigastria.
Barbilophozia kunzeana f. acuta (Schust.)
Potemk. comb. nov. (Lophozia kunzeana f.
acuta Schust. Hep. Anth. N. Am. 2:299,
1969)
NT:
Neromayaha I.
In dwarf shrub moss-lichen tundra, in
Polytrichum-lichen. mat.
Barbilophozia kunzeana f. rotundiloba
(Schust.) Potemk. comb. nov. (Lophozia kun-
zeana var. rotundiloba Schust., in Schuster &
al.,
Natl. Mus. Canada Bull. 164:24, 1959)
AT:
Kharasvey I.
In moss-sedge bog, among Sphagnum.
Plants green, very large, 10 - 15 mm long
and 1-1.5 mm wide, with oblique inserted,
distinctly dorsally decurrent 2 - 3 - lobed
wide leaves (2 - lobed leaves 840 - 1340
mkm wide and 730 - 1000 mkm long; 3 -
lobed leaves 1060 - 1400 mkm wide and
810 - 950 mkm long), near ventral and occa-
sionally dorsal base with 1-2 teeth or short
cilia; sinuses acute, gamma-like, with
strongly reflexed margins, descending 0.37 -
0.50 to 0.57 - 0.62(0.64); lobes mostly more
or less divergent, wide ovate, rotund, obtuse,
rarely acute; underleaves very large, 500 -
600(890) mkm long, bifid almost to the
base,
often with 1-2 cilia on outer margins
and, occasionally, 1 cilium per inner margin;
underleaf sinus often reflexed; marginal leaf
cells 14-20 mkm, median - 20 -
25(28) x 19 - 23 mkm; cuticle from faintly to
distinctly striolate-papillose, occasionally
with rather large flat papillae.
Yamal plants of f. rotundiloba differ from
American ones in the more lax leaf position
and predominantly bilobed very large un-
derleaves with cilia on outer and inner mar-
gins.
In the deeply bifid leaves, large bifid
underleaves and sporadically coarsely papil-
lose cuticle these plants resemble Barbi-
lophozia quadriloba, differing in the wide
ovate lobes with mostly obtuse apex, pure
green color and larger size. On the other
hand they may be confused with B. floerkei
(F.
Web. & Mohr) Loeske, which has the
same pigmentation, size, very large under-
leaves with cilia on inner margins of lobes
and gibbous leaf sinuses. B. kunzeana f. ro-
tundiloba is distinguished from this species
in the predominantly bilobed leaves with
considerably deeper sinuses and mostly
obtuse lobes.
Barbilophozia kunzeana f. wenzelioides
(Schust.) Potemk. [Novosti Sist. Nizsh. Rast.
29:167,
1993]
NT:
Bovanenkovo 1.
In weakly polygonal dwarf shrub lichen-
moss tundra, among Sphagnum. With gem-
mae.
Barbilophozia binsteadii (Kaal.) Loeske
(Orthocaulis attenuatus auct. non (Mart.)
Evans - Andrejeva 1981; Zhukova & Rebris-
taya 1986)
AT IV - V; NT IV - V; ST V.
In diverse wet moss tundras and bogs, usu-
ally among Sphagnum, Dicranum, Aulacom-
nium,
Polytrichum, considerably more rarely
among lichens, sporadically individual shoots
on bare soil. Occasionally with gemmae
which, as a rule, develop on plants of mod.
densifolia-colorata, only once on plants of
mod. laxifolia-viridis. Male and perianth
bearing plants occur rarely.
The species is very malleable ecologically
and morphologically. Occasionally it resem-
bles very much in some important diagnostic
features Barbilophozia attenuata (Mart.)
Loeske and B. atlantica. In such cases only
complex analysis of all diagnostic criteria
makes it possible to positively identify the
species (cf. Schuster 1969).
In the course of investigation of the species
in Yamal some paradoxical manifestations of
its variability were found: 1) not only neigh-
bour plants (Schuster 1969) but different
leaves of the same shoot may have cell net of
Barbilophozia attenuata- and B. binsteadii-
types,
i.e. with marginal cells (13)14-

The Hepaticae of the Yamal Peninsula 65
17(18) mkm and median 16 - 18 x 17 - 23
mkm with small concave-sided trigones or
with marginal cells 19-21 mkm and median
- 18 - 20 x 20 - 23 mkm with large nodulose
often confluent in lobes trigones; 2) leaves
are not always with 3 narrow lobes, occa-
sionally the plants with subtransverse in-
serted wide 3(4) - lobed B. atlantica-like
leaves with shallow sinuses (to 0.35 - 0.25 of
the length) occur. They differ from B. at-
lantica in smaller cells with often confluent
trigones in lobes and in the absence of un-
derleaves.
In this species, as in some others
(Anastrophyllum minutum, Barbilophozia
hatcheri, Cephalozia bicuspidata, etc.), the
plants affected by small nematodae ( 15-25
mkm in diam. and 300 - 1000 mkm long)
were found. Their apical leaves become
coarser, develop unusual for the species pur-
plish, blackish fuscous, or blackish purple
pigmentation and form the characteristic
"head" covering the nematodae.
Barbilophozia atlantica (Kaal.) K. Muell.
NT: Tomboitosyo I; ST: Khutyyaha I.
In dwarf shrub-sedge tundra and Sphag-
num willow stand in flood plain, associated
with Lophozia ventricosa, L. incisa and Pti-
lidium ciliare.
The plants from Tomboitosyo are mostly
quite typical. They are characterized by wide
leaves (width/length ratio of 3-lobed leaves -
1.42 -
1.67:1)
divided in 0.25 - 0.45 their
length in (2)3 wide obtuse to apicuiate (with
1 - celled apiculus) lobes; large cells,
marginal - 21-26 mkm, median - 25 -
32(35) x (21)24 - 28 mkm, with weakly
convex, never confluent trigones; underleaves
near shoot apexes; sporadic occurrence of
stalked slime papillae near the base of ven-
tral leaf margin. Together with plants char-
acterized above, several shoots with predomi-
nantly bilobed leaves were mentioned. The
distinctive feature of the plants from
Khutyyaha is an absence of amphigastria. In
the other respects they are quite typical. All
Yamal plants of Barbilophozia atlantica have
gamma-like, somewhat reflexed sinuses, giv-
ing them a peculiar appearance.
Subgen. BARBILOPHOZIA
Barbilophozia hatcheri (Evans) Loeske
AT: Tambey III, Kharasavey III; NT: Matyuiyaha I,
Tiutey 1, Neromayaha I, Mantyto II, Marre Sale 6, Se-
bayaha /, Khakhayayaha II, Lyakkotosyo П; ST:
Khutyyaha II, Khevesyo I.
In diverse herb communities on sandy and
loamy sand soils situated mostly on steep,
occasionally nival, slopes; in diverse lichen
tundras on sandy soil. Almost always with
gemmae.
Together with quite typical plants the
plants of mod. parvifotia-colorata-eciliata-
gemmipara occur often. They have small size,
to 2 - 4 mm long and 0,5 mm wide, deep
brown pigmentation, (2)3 - lobed or pre-
dominantly bilobed leaves with obtuse or
briefly apicuiate lobes and mostly without
cilia near ventral leaf base (only on individ-
ual leaves solitary, short cilia composed of
2-4 cells can be found). The plants of this
modification may be confused with small
forms of Barbilophozia barbata. They differ
from it, however, in considerably smaller
size,
sporadic occurrence of cilia near ventral
leaf base, underleaf structure, and constant
presence of gemmae.
Occasionally Barbilophozia hatcheri is asso-
ciated with B. lycopodioides. In Lyakkotosyo
transitional forms of the both species from
mod. parvifolia-colorata to mod. megafolia-
viridis were found in one specimen. In this
case B. hatcheri differed from B. lycopodi-
oides in constant presence of gemmae (even
small plants of mod. parvifolia-colorata of B.
lycopodioides with 2 - 3 - lobed leaves do not
develop gemmae), shorter cilia and lobe api-
culi composed of shorter cells and almost
straight, not distinctly convex lobe sides.
These observations confirm the taxonomic
status of B. hatcheri as a species, not a vari-
ety of B, lycopodioides (cf. Schljakov 1980).
Barbilophozia rubescens (Schust. &
Damsh.) Karttunen & Soederstroem {B.
hatcheri var. grandiretis Lammes)
AT: Tambey I, Kharasavey SI; NT: Bovanenkovo /,
Marre Sale /, Khakhayayaha II,
In moss tundras and herb communities on
nival slopes, associated with Ptilidium ciliare,
Barbilophozia hatcheri, B. lycopodioides and
Tritomaria quinquedentata. Once

66 A. D. POTEMKIN
(Kharasavey) with few gemmae, 25 -
36 x 23 - 34 mkm, deep purple or reddish
brown, mostly 2 - celled, 3 - 4 - angular to
polygonal, with feebly projecting angles (cf.
Schuster 1988).
Barbilophozia rubescens, for comparison
with B. hatcheri and B. lycopodioides is char-
acterized by robust size contrasting with usu-
ally small underleaves occurring only near
shoot apicies; fewer cilia (1-3) sporadically
developing near ventral leaf base and com-
posed of not strongly elongated rectangular
cells (30 - 60 x 14 - 20 mkm, width/length
ratio 1:1.5 - 3.0), larger cells with numerous
oil-bodies (5)8 - 16(18) per cell, etc.(see also
Schuster 1988).
Barbilophozia lycopodioides (Wallr.)
Loeske
AT: Kharasavey I; NT: Matyuiyaha I, Marre Sale 3,
Saletayaha 3, Khakhayayaha III, Lyakkotosyo I; ST:
Khutyyaha I, Yuribey 2.
In herb communities at the foot of steep
slopes and wet herb-grass tundra, associated
with Barbilophozia hatcheri, B. rubescens,
Tritomaria quinquedentata, Ptilidium ciliare,
and Lophozia heterocolpos. Almost always
without gemmae, occasionally with perianths.
Barbilophozia barbata (Schmid. ex Schreb.)
Loeske var. barbata
NT: Matyuiyaha I, Bovanenkovo /, Neromayaha I,
Marre Sale I, Lyakkotosyo I.
In moss willow stands and lichen-moss
tundras.
Barbilophozia barbata var. amphigastriata
K. Muell.
NT: Neromayaha I, Marre Sale 1.
In herb-willow-moss tundra and flood plain
willow stand, with B. barbata var. barbata
and Ptilidium ciliare.
The varietal status of var. amphigastriata is
doubtful. Possibly it is only a modification of
the polymorphous Barbilophozia barbata.
LOPHOZIA
(Dum.) Dum.
Subgen. PROTOLOPHOZIA Schust.
Lophozia debiliformis Schust. & Damsh.
NT: Khakhayayaha I, Sebayaha 1; ST: Er'yaha I.
In willow stands on slope, late snow area
and bank of brook, usually on fine-grained
soil with Tritomaria quinquedentata f. gra-
cilis, Lophozia excisa, L. ventricosa s.l.,
Scapania curta var. grandiretis, Barbilophozia
hyperborea, B. lycopodioides and Pleuro-
cladula albescens, or in pure mats. In
Khakhayayaha with gemmae.
Subgen. LEIOCOLEA K. Muell.
The species of Leiocolea are basiphilous
and, consequently, very rare in Yamal oc-
curing only in sites with comparatively rich
soils.
The only acid-tolerant species of the
subgenus, Lophozia heterocolpos, is not un-
common on the peninsula.
Lophozia rutheana (Limpr.) M.A. Howe
NT: Bovanenkovo 1.
In cotton grass-sedge-hypnum moss bog,
among Calliergon sarmentosum, Scapania ir-
rigua var. rufescens, Barbilophozia quadriloba
f. glareosa, Tritomaria quinquedentata, etc.
Lophozia gillmanii (Aust.) Schust.
NT: Bovanenkovo 2; ST: Khutyyaha I.
In Equisetum and Sphagnum willow stands
with Plagiochila asplenioides subsp. porel-
loides, Aneura pinguis, Lophozia heterocolpos
var. arctica, Chiloscyphus fragilis, Mesopty-
chia, Blepharostoma, etc.
Lophozia collaris (Nees) Dum.
AT: Belyy I (Zhukova & Rebristaya 1987); ST: Lap-
tayaha I.
In spotty herb-willow Racomitrium - Gym-
nomitrion tundra and in tundra shortgrass
meadow; individual plants and small tufts in
association with Nardia geoscyphus,
Cephalozia pleniceps, and Lophozia sp.
Lophozia heterocolpos (Thed. ex Hartm.)
M.A. Howe var. heterocolpos
AT: Belyy (Arnell 1918), Khabeiyaha I, Kharasavey I;
NT: Matyuiyaha I, Tiutey 2, Bovanenkovo /,
Tomboitosyo I, Neromayaha II, Saletayaha J, Yuribeitoy-
aha IV, Lyakkotosyo I; ST: Khevesyo I; Laptayaha II.
In diverse wet tundras, occasionally in late
snow areas, once in fire place. Often with
gemmae, once with perianths.
Lophozia heterocolpos var. arctica (S.
Arn.) Schust. & Damsh.
AT: Kharasavey I; NT: Neromayaha III; Yuribeitoyaha
П, Khakhayayaha I, Lyakkotosyo I; ST: Khutyyaha I,
Khevesyo I.
In Equisetum-herb, herb-dwarf shrub-moss
and willow-sedge-moss tundras, at the foot of

The Hepaticae of the Yamal Peninsula 67
steep slopes, associated with Blepharostoma
trichophyllum var. brevirete, Lophozia hetero-
colpos var. heterocolpos, L. excisa, Ptilidium
ciliare, Tritomaria quinquedentata, T. scitula,
Barbilophozia spp., Odontoschisma macounii,
Scapania cuspiduligera, Preissia, etc. Spo-
radically with gemmae and perianths. The
gemmae often develop on rather stout flagel-
lae with reduced leaves, similar to those of
var. heterocolpos. The perianth in var. arctica
lacks a beak and is more or less plicate, at
least with one plica in dorsal part when im-
mature and smooth when mature (only one
mature perianth has been seen). The mouth
of the perianth varies from faintly crenulate
to sparsely dentate and the teeth are com-
posed of 1 - 3 subisodiametric cells.
The degree of manifestation of the perianth
beak in the type variety of the species varies
greatly - from undetectable or rudimentary
to rather long and narrow. A plicate perianth
surface occurs in var. heterocolpos also (cf.
Schuster 1974, Fig. 163: 8, 9). Consequently,
the presence of beak and plicae considered as
significant generic or infrageneric criteria (cf.
Mueller 1951 - 1958; Inoue 1957, 1961;
Schuster 1974; Schljakov 1980, etc.) has only
a little infraspecific value.
This comparatively poorly known taxon is
rather common, not only in Yamal but in the
Asian Arctic as a whole. It differs from the
type variety of the species by larger cells
(marginal - 20 - 30(34) x 23 - 35(40) mkm,
median - 23 - 35 x 24 - 42 mkm) and gem-
mae (18 - 23 x 24 - 38 mkm), a mostly
smooth or faintly papillose cuticle, more nu-
merous oil-bodies ((3)4 - 11(16) per cell),
Barbilophozia barbata-like leaves that are
2(3) - lobed, exceptionally 4 - lobed, often
developing purplish pigmentation, etc.
Lophozia heterocolpos var. harpanthoides
(Bryhn & Kaal.) Schust.
AT: Belyy II.
In herb-willow-moss tundras with Ble-
pharostoma trichophyllum var. brevirete, Trit-
omaria quinquedentata, Odontoschisma ma-
counii, Scapania zemljae, Jungermannia po-
laris, Cephaloziella arctica, etc.
The investigated plants are mod. meso-
derma-colorata of the variety with straight-
sided, rarely weakly convex, never confluent
trigones. Small-leaved phenotypes often de-
velop purplish violet pigmentation in distal
parts of leaves. One shoot with a ventral-in-
tercalary branch originating from the under-
leaf axil was found also. Otherwise the mate-
rial is quite typical. It has narrow and shal-
low sinuses with convex sides, usually rotund
lobes,
often bilobed amphigastria, and com-
paratively small, subisodiametric basal and
median leaf cells (median ca. 20 - 31 x 17 -
24 mkm).
Lophozia badensis (Gott. ex Rabenh.)
Schiffn. ex Dalla Torre & Sarnth.
AT: Kharasavey I; NT: Yuribeitoyaha I.
In hillock tundra on brooklet bank and in
dwarf shrub spotty tundra on bare soil, asso-
ciated with Jungermannia polaris, Arnellia
fennica, Scapania gymnostomophila f. in-
curva, Tritomaria scitula, Lophozia heterocol-
pos,
and Barbilophozia quadriloba f. glareosa.
In Kharasavey plants of mod. parvifolia-
mesoderma-colorata were found. The small-
est plants of this modification have subtrans-
versely oriented, more or less concave and
fuscous leaves and resemble small species of
Marsupella. In the sterile state they differ
from Marsupella by stem anatomy and com-
paratively large cells.
Subgen. SCHISTOCHILOPSIS Kitag.
Lophozia grandiretis (Lindb. ex Kaal.)
Schiffn.
ST: Khevesyo III, Laptayaha II, Er'yaha I.
On spots in spotty moss and lichen-moss
tundras, bare loamy soil near temporary wa-
ter course and peat outcrop, associated with
Cephalozia pleniceps, Blepharostoma tri-
chophyllum var. brevirete, Anthelia ju-
ratzkana, Nardia geoscyphus, Tritomaria het-
erophylla, T. quinquedentata, Aneura pinguis,
Odontoschisma macounii, etc. Gemmae usu-
ally develop in small quantity. Male plants
were found once. Antheridia vary from green
to yellow and purplish. The stalks are 2 - or,
at least partly, 3 - seriate.
The color varies from green to deep purple.
Plasmolysis, resembling that of Lophozia
opacifolia, occurs exceptionally in plants of
mod. viridis. Such plants differ from L.

68 A. D. POTEMKIN
opacifolia in their dirty green, not bluish
green coloration, as well as the size of cells
and gemmae.
Lopozia incisa (Schrad.) Dum. s.sir.,
AT: Tambey i, Khabelyaha II, Kharasavey H: NT:
Matyuiyaha I, Bovanenkovo J, Tombostcsyo J, Nero-
mayaha Ш, Mantyio I, Saietayaha /, Yuribeitoyaha I,
Khakhayayaha II, Lyakkotosyo I; ST: Khutyyaha II,
Khevesyo Ш, Laptoyaha I, Er'yaha I, Khadytayaha /
In diverse wet moss tundras, bogs, peat
outcrops, often among Sphagnum. Usually
with gemmae. Male plants were found only
once.
Among Sphagum the plants of mod. laxifo-
lia-integrifolia, resembling Lophozia laxa
(Lindb.) Grolle, sporadically occur. They
differ from this species by opaque bluish
green coloration, absence of purplish pig-
ment, character of gemmae, etc. Moreover L.
laxa is an amphi-oceanic species apparently
unknown from continental regions of Eurasia
(Konstantinova & al., 1992).
When sterile differentiation of L. incisa
from L. opacifolia is problematic on account
of considerable overlapping of their variabil-
ity (cf. Schuster 1969, 1988, etc.). Usually
plants of L, incisa are smaller, with thinner
stems,
and narrower, deeper divided leaves,
2 - 3 - stratose only near the base.
Moreover, L. incisa is associated with
acidophilus species, while L. opacifolia
occurs on enriched soils.
Lophozia incisa var. inermis K. Muell.
AT: Tirvyyaha 1.
In wet willow-moss tundra among Sphag-
num sp., Barbilophozia binsteadii, Ptilidium
ciliare, Blepharostoma trichophyllum var,
brevirete. In my opinion, the most character-
istic feature of this rare variety, which is un-
known for the other taxa of the subgenus, is
the prominent bulging trigones.
Lophozia opacifolia Culm, ex Meyl.
AT: Belyy II, Khabeiyaha III, Tambey III, Kharasavey
II;
NT: Matyuiyaha I, Bovanenkovo /, Tomboitosyo I,
Neroaaygha III, Mantyio I, Yuribeitoyaha II,
Khakhayayaha II; ST: Khutyyaha I, Khevesyo III,
Yiiribcy 2, Er'yaha I.
On spots of bare soils in diverse tundras,
late snow areas at the foot of slopes and peat
outcrops, and on brooklet banks. Usually
with gemmae. Maie plants and plants with
perianths and mature capsules rare.
Some phenotypes of Lophozia opacifolia re-
semble L. hyperarctica Schust. They have
very thick, fleshy and flattened, elliptical in
cross section, blackish brown pigmented with
age stems; predominantly bilobed, very wide,
almost transversely inserted dorsally leaves,
with shallow crescentic sinus and entire or
faintly denticulate (on gemmiparous shoots)
margins; and large gemmae, 20 -
34(40) x 20 - 31 mkm. They differ from L.
hyperarctica in opaque, whitish green col-
oration; characteristically plasmolysed cell
contents; absence of black pigmentation of
cell walls in old parts of ventral sector of
stem (brownish black coloration of old part of
stem of L. opacifolia connected with destruc-
tion of the ceil walls, not with black pigmen-
tation);
presence of diffuse mycorrhiza in
ventral part of stem.
Subgen. ISOPACHES (Buch) Schust.
For a description of Lophozia alboviridis
and L decolorans in Yamal and differentia-
tion of the species of the subgenus see
Potemkin (1990c). I would like to note here
only the most important distinctive criteria of
the species of the subgenus in the key given
below and to stress that sex distribution is
not constant in arctic populations of
Isopaches species: L. alboviridis in Yamal, as
well as in Greenland (Schuster & Damsholt
1974) is mostly dioicous but sporadically
paroicous; L. decolorans in Yamal is paro-
icous,
but in the other localities dioicous; in
L bicrenata male plants occur sporadically,
while as a rule this species is paroicous.
KEY
TO SPECIES OF ISOPACHES
1.
Marginal leaf cells considerably smaller
than median, 11 - 18(20) mkm; leaf lobes
more or less unequal, ventral lobe longer and
wider than dorsal, ending in sharp tips
formed by (1)2-3(4) superimposed cells
with distinctly elongated apical cell; gemmae
with strongly projected and very thick angles,
mostly regulary stellate, 19 - 25(26) x 16 -
24 mkm L alboviridis
1.
Marginal cells not distinctly different
from median, larger than 20 mkm; leaf lobes
usually almost equal, ending in obtuse to

The Hepaticae of the Yamal Peninsula 69
sharp apiculi formed by 1(2) superimposed
subisodiametric cells; gemmae variable, from
mostly irregularly polygonal to rectangular
and triangular, with feebly to rather strongly
projecting and thickened angles, as an ex-
ception - oval, 17 - 25 x 23 - 31 mkm (Loph-
ozia bicrenata) or 17 - 30(34) x 23 -45(56)
mkm (L. decolorans) 2
2.
Leaves usually densely imbricate, wide
(width/length ratio usually considerably
more than
1:1.2),
trapezium-like, with mostly
shallow wide crescentic sinus, 0.05 - 0.15 of
the leaf length (sterile plants resemble Pras-
anthus suecicus); male bracts with antical
tooth; perianth mouth shortly ciliate, cilia 1 -
2(3) - celled; often with stolons.
L.
decolorans
2.
Leaves not densely imbricate, their width
not obviously exceeding their length
(width/legth ratio ca. 1:0.9 - 1.2); sinus
deeper, 0.2 - 0.3 of the leaf length, mostly
U-shaped, occasionally crescentic (plants es-
sentially different from Prasanthus suecicus);
male bracts usually without antical tooth; pe-
rianth mouth spinose-ciliate, cilia 2 - 4 -
celled; stolons absent L. bicrenata
Working with this key, the following corre-
lation should be taken into account: gemmae
are more angulate and thick-angled but cilia
of perianth mouth and apiculi of leaf lobes
are shorter and leaves are denser in plants of
exposed sites and just the opposite for plants
of shade sites.
Lophozia bicrenata (Schmid. ex Hoffm.)
Dum.
ST:
Khutyyaha I.
On thinly turfed sandy soil in dwarf shrub-
herb moss tundra on steep slope, in
association with L. alboviridis, Gymnomitrion
corallioides, Lophozia excisa and
Cephaloziella arctica. With perianths and
gemmae.
Lophozia alboviridis Schust.
AT:
Kharasavey I; NT: Matyuiyaha I, Neromayaha III,
Mantyto
I, Saletayaha 2, Khakhayayaha II, Lyakkotosyo
I;
ST: Khutyyaha III, Khevesyo I, Laptayaha I, Er'yaha
I.
On weakly turfed sandy and loamy sand
soils,
sometimes with the other species of the
subgenus. Always with gemmae; male plants
and plants with perianths occur rarely; seen
once with mature capsules. Potemkin (1990c,
fig.
1) provides illustrations of this species by
material from Yamal.
Lophozia decolorans (Limpr.) Steph.
AT:
Tambey II; NT: Lyakkotosyo I; ST: Khutyyaha I.
On steep herb sandy slopes, associated with
Barbilophozia hatcheri, Gymnomitrion coral-
lioides, Lophozia excisa, L. alboviridis,
Scapania praetervisa, Anastrophyllum minu-
tum,
etc. With gemmae and perianths.
Potemkin (1990c, fig. 2) provides illustra-
tions of this species by material from Yamal.
Subgen. LOPHOZIA
Lophozia heteromorpha Schust. & Oamsh.
(Fig. 3)
NT:
Bovanenkovo /, Neromayaha I; ST: Khevesyo I.
In spotty dwarf shrub formation with
Betula nana, among Dicranum elongatum
with Lophozia excisa var. infuscata, L. major,
L.
heterocolpos, Blepharostoma trichophyllum
var. trichophyllum; in lichen-moss-dwarf
shrub tundra, among Sphagnum with Barbi-
lophozia kunzeana; at the foot of steep herb
slope near margin of sedge bog with Pellia
neesiana, Chiloscyphus fragilis, Scapania ir-
rigua, Cephalozia pleniceps, and Jungerman-
nia hyalina. With gemmae.
Lophozia heteromorpha is a polymorphous
species with unclear limits of variability. The
most important features, differentiating it
from the other taxa of Lophozia, are: po-
lymorphous leaf form; violetish pigmentation
of mature gemmae; violet pigmentation of
young leaf margins in distal parts, replaced
by fuscous pigment with age; frequent
presence of amphiga stria; and weak
dorsiventral differentiation of stem tissue. All
these characters are not constant, however,
and only a complex analysis of them all per-
mits sure identification of the species (see
also Potemkin 1990a).
Lophozia jurensis Meyl. ex K. Muell. (L.
latifolia Schust.; L. sudetica auct. non (Nees
ex Hueb.) Grolle p.p. - Potemkin 1989; L.
pellucida var. minor auct. non Schust. -
Potemkin 1988, 1989)
AT:
Belyy II, Tambey II, Kharasavey II; NT:
Matyuiyaha
II, Tiutey 2, Bovanenkovo 1, Neromayaha
III,
Yuribeitoyaha I, Khakhayayaha I, Lyakkotosyo II;
ST:
Khutyyaha I, Khevesyo II, Laptayaha II, Kharangy-
neto
4, Er'yaha I, Khadytayaha 3.

70 A. D. POTEMKIN
200 дш
100 цт
Fig. 3. Lophozia
heteromorpha.
1 - Sterile shoot; 2, 3 - Gemmae; 4 - Median cells with oil-bodies; 5-7 - Underleaves;
8 - Magnified underleaf from Fig. 7; 9-14 - Leaves; 15 - Cross section of stem. Scale bars: 1 mm - for 1; 0.5 mm - for
5 - 7, 9 - 14; 200 mkm - for 8; 100 mkm - for 15; 50 mkm - for 2 - 4. All from Khevesyo (After Potemkin 1990a).
In diverse moss tundras. Often with peri-
anths,
rarely with gemmae and mature cap-
sules.
Yamal plants are mostly paroicous. Individ-
ual monoicous shoots were found only in
Yuribeitoyaha, Lyakkotosyo and Laptayaha.
The species varies from mod. laxifolia-viridis
to mod. densifolia-coloraia. Secondary pig-
mentation is usually purplish red. Plants of
mod. laxifolia-viridis often represent pheno-
copies of Lophozia pellucida var. minor (cf.
Schuster 1969, Fig. 220:11). They differ
from this species in: 1) almost constant
absence of gemmae (if gemmae present, they
are reddish brown or reddish, with faintly
projecting angles (vs strongly angulate,
colourless to yellow and yellowish brown
gemmae of L. pellucida); 2) male bracts with
antical teeth; 3) ecological behaviour - L.
jurensis associated with acidophilous species
while L. pellucida with basiphilous ones.
Lophozia excisa (Dicks.) Dum. (? L unci-
nata Schljak. syn. nov.).
In Yamal three varieties of L. excisa, repre-
senting apparently more or less distinct enti-
ties of this genetically highly complicated
species are distinguished. My approaches to
their differentiation are given in the key.
KEY
TO YAMAL VARIETIES OF LOPHOZIA
EXCISA
1.
Perianth mouth crenulate or dentate,
composed of more or less regulary projecting,
oriented perpendicularly to margin, elon-

The Hepaticae of the Yamal Peninsula 71
gated, finger-like cells. Mature gemmae red-
dish brown to dull purplish 2
1.
Perianth mouth with smooth margin,
composed of mostly tangentially oriented
cells,
and scattered 1 - 2(3) - celled firm
teeth formed by mostly subisodiametric cells.
Mature gemmae colourless to violetish
L.
excisa cf.var. succulenta
2.
Perianth mouth crenulate; marginal cells,
in lobes, 17 - 30 mkm (tangentially mea-
sured) L. excisa var.
excisa
2.
Perianth mouth dentate, with free teeth
0.5 - 2(3 - 4) cells long; marginal cells, in
lobes,
(25)30 - 40(42) mkm (tangentially
measured) L. excisa.var. infuscata
Lophozia excisa var. excisa (L. sudetica
auct. non (Nees ex Hueb.) Grolle p.p. -
Potemkin 1989)
AT:
Belyy II, Khabeyyaha HI, Tambey III, Kharasavey
II;
NT: Matyiyaha I, Tiutey 1, Bovanenkovo .',
Tomboitosyo II, Neromayaha IV, Mantyto I, Marre Sale
/, Saletayaha 7, Yuribeitoyaha II, Khakhayayaha III,
Lyakkotosyo III; ST: Khutyyaha HI, Khevesyo III,
Yuribey 1, Laptayaha II, Kharangyneto 1, Er'yaha II.
In diverse moss and spotty tundras, on
herb slopes, late snow areas, peat outcrops,
moss bogs, banks of water courses and lakes.
Often with perianths and gemmae, sporadi-
cally with mature capsules.
An euritopic, extremely malleable taxon,
varying from mod. laxifolia-viridis-egemmi-
para, occurring in wet sites among mosses, to
mod. parvifolia-densifolia-colorata-gemmi-
para, which is characteristic for habitats on
bare soil. The smallest plants of the last
modification are only 0.8 - 0.9 mm long and
0.2 - 0.35 mm wide. They usually have a
thick, fleshy stem and comparatively small
leaves as well as comparatively small cells,
(17)22 - 30 x (17)20 - 25(28) mkm, with
small trigones. Their gemmae are 2 - celled,
21 - 31(36) x 17 - 25(28) mkm, and from
brown and brownish red to deep purple.
Yamal plants of Lophozia excisa var. excisa
develop brown pigmentation more often than
red or purplish. This may lead to confusing
L.
excisa and L. major. Such forms of L. ex-
cise, however, have leaves not so strongly
narrowed in upper part, as in L. major, with
the maximal width between the middle and
lower thirds or in the middle, not near the
base,
and almost triangular, not horn-like
lobes (if the maximal width of leaves of L.
major is in the middle, their ventral margin
is very convex but the dorsal is almost
straight, i.e. the leaves of this species are
usually considerably more asymmetric than
in L, excisa).
Small gemmiparous plants of Lophozia ex-
cisa are confused also with L. sudetica. They
differ from this species in their wide ventral
merophytes, unmodified gemmiparous leaves,
also often in larger cells as well as size,
shape and coloration of gemmae, etc. More-
over L. excisa differs from all other regional
species of the subgenus, excepting L. juren-
sis,
in paroecia and, as a consequence, in
often developing perianths.
•a
Lophozia excisa var. infuscata Schust &
Damsh.
AT:
Khabeiyaha I; NT: Neromayaha II, Lyakkotosyo I;
ST:
Er'yaha I.
In willow stands on humused soil with Bar-
bilophozia binsteadii, B. kunzeana, B.
quadriloba, Anastrophyllum minutum,
Cephalozia pleniceps, Tritomaria quinque-
deniata, Blepharostoma, etc.; in wet moss
tundra among Aulacomnium palustre, with
Tritomaria quinquedentata and Lophozia
major. With gemmae, perianths and mature
capsules.
The differentiation of this variety, recently
described by Schuster and Damsholt (1974)
and known before this report only from
Greenland, is sometimes rather problematic.
Its distinctive criteria, mentioned by the au-
thors - larger cells (marginal in lobes - (26 -
29)30 - 38(40) mkm, median - (32 - 35)36 -
45(48) x (36)40 - 55(60) mkm) and gem-
mae,
(28)30 37 x 33 - 38(48) mkm; more
numerous oil-bodies, (12 - 18)20 - 36(40 -
45) per cell; fuscous secondary pigmentation;
dentate perianth mouth with free teeth 0.5 -
2 (3 - 4) cells long and terminal cells (where
free) ca. 56 - 68(75 - 80) x 16.5 - 22 mkm -
are mainly quantitative. Their variability
overlaps with that of the type variety. The
most reliable of these criteria, for separation
of the Yamal plants of the variety, are
character of perianth mouth and size of
marginal cells of leaf lobes.

72 A. D. POTEMKIN
Lophozia excisa cf. var. succulenta Schust.
& Damsh,
AT:
Belyy П, Khabeiyaha II, Tambey I, Kharasavey I;
NT:
Neromayaha I; ST: Khutyyaha I, Khevesyo I.
On bare fine-grained soil in late snow areas
and diverse spotty tundras; mostly individual
shoots among Prasanthus suecicus, Gym-
nomitrion corallioides, G. concinnatum, Nar-
dia geoscyphus, Scapania scandica, Trito-
maria quinquedentata f. gracilis, Lophozia
ventricosa s.l., Cephalozia bicuspidata, An-
thelia juratzkana, Cephaloziella arctica,
Anastrophyllum minutum, Jungermannia
sphaerocarpa var. nana, etc. Often with peri-
anths,
gemmae and mature capsules.
The variety was described from West
Greenland by Schuster and Damsholt (1974)
on the base of "sparing type material",
which was unavailable for critical study. It is
obvious that this material can not reflect the
phenetic variability of the taxon completely.
It gives me the basis to attribute to the cf.
var. succulenta the Yamal plants that differ
somewhat from the type until an
investigation of the type is possible. Because
of the tentative identification of these plants,
their description and differences from the
type are listed.
Plants minute to small, 1-10 mm long and
(0.2)0.5 - 1.1 mm wide, resembling small
compact soil forms of Lophozia excisa var.
excisa. Stem more or less fleshy, thick, con-
spicuously mycorrhizal throughout much of
the medulla. Leaves usually subrotund,
broader than long, imbricate, subvertical,
often 2(3) - stratose at base; sinus mostly
crescentic, occasionally gibbous, ca. 1/5 -
2/5 of the leaf length; lobes acute; distal
parts,
especially margins, of young leaves
bright red to deep purple and violet, of ma-
ture and old leaves - fuscous (brown pigment
replaces red pigment with age), unpigmented
parts of leaves and stem in dead material
opalescent. Cells of leaves thin-walled, except
for small to sporadically rather pronounced
trigones; marginal cells in lobes, 17-28
mkm (measured tangentially), median from
14 - 17 x 17 - 23 mkm to 20 - 25(28) x 25 -
30(35) mkm; oil-bodies small, 3 - 5(6) mkm,
4 - 12(14) per cell, according to Schuster and
Damsholt (I.c.) occasionally 18-20 per cell.
Gemmae green to violetish (only individual
"overmatured" gemmae deep violet), 1-2
celled, usually thin-walled, almost oval to
triangular and polygonal, with weakly
projected angles, 20 - 23 x 23 - 34 mkm.
Paroicous. Male bracts leaf-like, often with
somewhat gibbous base, 1 - 2 - androus,
with or without distinct antical tooth. Female
bracts 2 - 3 - fid; lobes acute to apiculate
(apiculus 1 - 3(5) cells long) often with
strongly reflexed margins; sinuses rather
deep,
angulate, gibbous. Perianth 2-5(10) -
stratose at base, usually bistratose up to
middle or higher, at mouth usually bleached,
weakly lobulate, occasionally formed of
rather thick-walled collenchymatous cells; the
mouth margin smooth, composed of mostly
tangentially oriented cells, with scattered 1 -
2(3) - celled firm teeth formed by mostly
subisodiametric, only as an exception by
elongated cells. Capsule wall 3 - 4 - stratose
with epidermal layer usually considerably
thicker than any of inner ones (the thick-
nesses ratio ca. 1.5 - 2.0:1). Spores 16 -
18(20) mkm. Elaters 2 - spiral, (7)8-9(11)
mkm thick.
In their description of this variety Schuster
and Damsholt (I.c.) stressed that "the com-
bination of denticulate, firm-celled perianth
mouth, the polystratose perianth base, and
edentate bases of male bracts isolate these
plants from any other variety of L. excisa".
In my opinion, the most important of these
criteria is the character of perianth mouth -
the combination of scattered teeth and
smooth margin, composed of tangentially
elongated cells (a firm-celled peranth mouth
is not constant for the Yamal plants of var.
succulenta and it has been seen in var. infus-
cata also). The character of the perianth
base as well as the leaf base (the authors
note unistratose leaf base for the variety) is
not a very reliable feature. According to my
per-sonal observations a 2 - 3 - stratose
perianth base is not a rarity for the typical
variety of the species. A bistratose leaf base
and edentate male bracts occur exceptionally
in var. excisa also. On the other hand,
"sparing type material" cannot show the
degree of constancy of these characters and
the constancy of the absence of gemmae

The Hepaticae of the Yamal Peninsula 73
stressed in the description.
Lophozia cf. longidens (Lindb.) Macoun
AT:
Khabelyaha II, Tambey I.
In lichen and lichen-moss tundras, individ-
ual plants among lichens with Anastrophyl-
lum minutum, Tritomaria quinquedentata,
Ptilidium ciliare, Gymnomitrion concinnatum,
etc.
With gemmae.
Because some species of Lophozia in certain
conditions may develop phenocopies of L.
longidens and collections of the species in
Yamal are sparing and unsufficient for its
sure identification it listed under the sign
"cf.".
Lophozia cf. savicziae Schljak. (L. wenzelii
auct. non (Nees) Steph. - Zhukova & Re-
bristaya 1986, 1987)
AT Ш; NT: Matyulyaha II, Bovanenkovo S,
Tomboltoeyo II, Neromayaha IV, Mantyto II,
Khakhayayaha II, Lyakkotosyo I; ST: Khutyyaha III,
Khevesyo V, Laptayaha II, Kharangyneto 2, Khady-
tayaha 4.
In diverse wet moss, moss-lichen and lichen
tundras, late snow areas, on spots in spotty
tundras, steep slopes and in bog communi-
ties.
Often with gemmae; male plants rare.
The most characteristic features of the
species are concave, trapezium-like, rather
rigid leaves, spread away from the stem from
their bases, with a rather shallow, usually
lunate sinus and mostly apiculate lobes; often
deep purple ventral sector of stem as well as
ventral leaf bases; polygonal to stellate gem-
mae with strongly thickened and projecting
angles; usually large cells.
In original description of the species Schl-
jakov (1973) noted two types of oil-bodies -
small, 2-4 mkm, homogenous, drop-like,
15 - 30 and more per cell, or, if they are
larger, they are less numerous, granulate,
with central droplet. Later (Schljakov 1980)
he listed granulate oil-bodies without central
droplet for the species also, interpreting the
polymorphism of oil-bodies as a feature of its
antiquity. Oil-bodies with central droplet, ca.
(5)8 - 18(20) to 15-30 per cell, 3 - 7 mkm
in diam., were found in most collections of
the species studied in a living state (mainly
from Bovanenkovo, Neromayaha, Khevesyo).
In some specimens of the species from the
same localities granulate oil-bodies without
central droplet were found. The differentia-
tion of (?) forms of the species with granu-
late oil-bodies and oil-bodies with central
droplet from the other large-celled Lophozia,
namely L. ventricosa var. grandiretis (Buch &
S. Arn.) Schust. & Damsh., L. ventricosa var.
rigida Schust. as well as L. wenzelii var. lap-
ponica Buch & S. Arn. sensu Schust. remains
unclear. As a consequence, the conception of
the species, particulary in respect of oil-body
polymorphism, remains unclear also.
Lophozia ventricosa (Dicks.) Dum. s.l.
(incl. L groenlandica (Nees) Macoun sensu
Schljak. 1975, non sensu Schust. 1969, excl.
syn. L heteromorpha Schust. & Damsh., )
AT:
Belyy I, Khabelyaha III, Tambey III, Kharasavey
II;
NT: Matyulyaha V, Tiutey 7, Venuieuo Л Bova-
nenkovo 4, Tomboltoeyo Ш, Neromayaha III, Mantyto П,
Marre Sale I, Saletayaha 1, Yuribeitoyaha II,
Khakhayayaha V, Lyakkotosyo HI; ST: Khutyyaha V,
Khevesyo V, Laptayaha III, Kharangyneto 4, Er'yaha III,
Khadytayaha 5.
In diverse moss and lichen tundras, herb
communities, late snow areas, moss bogs.
Usually with gemmae; perianth bearing and
male plants rare.
Lophozia ventricosa var. longiflora (Nees)
Macoun ( Z. longiflora (Nees) Schiffn. s.l.,
incl. L. porphyroleuca auct. non Schiffn.; L
ventricosa auct. non (Dicks.) Dum. p.p. -
Zhukova & Rebristaya 1986)
AT:
Belyy I, Khabelyaha ГУ, Tambey III, Kharasavey
II;
NT: Matyulyaha III, Tiutey 2, Venuieuo /, Bova-
nenkovo /, Neromayaha III, Mantyto V, Marre Sale 7,
Saletayaha 7, Yuribeitoyaha IV, Khakhayayaha П,
Lyakkotosyo IV; ST: Khutyyaha V, Khevesyo V, Yuribey
2,
Laptayaha V, Kharangyneto 5, Er'yaha III.
In diverse moss and lichen tundras, moss
bogs,
occasionally on spots in spotty tundras.
Perianth bearing and male plants rather
freq-uent; occasionally with gemmae .and
mature capsules.
Lophozia wenzelii (Nees) Steph.
AT:
Kharasavey I; NT: Matyulyaha I, Bovanenkovo I,
Neromayaha II, Sebayaha 1, Khakhayayaha I; ST:
Khutyyaha II, Khevesyo I, Laptayaha II, Er'yaha II.
In wet dwarf shrub-moss tundras, once in
herb-moss nival tundra. Perianth bearing,
male and gemmiparous plants occur sporadi-
cally.

74 A. D. POTEMKIN
Lophozia sudetica (Nees ex Hueb.) Grolle
(L.
rufescens auct. non Schljak. - Potemkin
1989)
AT: Tambey II, Kharasavey I; ST: Khutyyaha I, Lap-
tayaha I.
In diverse lichen tundras and spotty Gym-
nomitrion tundra, on steep slope; individual
shoots on fine-grained soil. With gemmae.
Lophozia sudetica is a rare species in Ya-
mal. It is represented by small forms like
"Lophozia alpestris var. gelida (Tayl.)
K.Muell." A part of the earlier reports of the
species from Yamal (Arnell 1918 (?);
Zhukova & Rebristaya 1986, 1987; Potemkin
1989) must be attributed to mod. parvifolia-
colorata-parviretis-gemmipara of L. excisa, L.
major and L. jurensis, which may develop
similar small cells and pigmentation. L.
sudetica differs from them, however, in con-
siderably more symmetrical leaves, which
usually become dentate from gemmae pro-
duction; narrow ventral merophyte and usu-
ally purplish pigmented ventral part of the
stem and ventral leaf base.
My attempts to use the leaf cell rows crite-
rion (Schljakov 1980) for separation of this
species and Lophozia rufescens Schljak. were
resultless. Other criteria for differentiation of
small forms of the both species with pig-
mented gemmae are absent. In my opinion
the cell row directions in specimens of L.
sudetica determined by Schljakov (LE) vary
principally from "in lobes" to "to sinus". I
am sceptical about this criterion for
differentiation of Lophozia species.
Lophozia major (C. Jens.) Schljak. (L. po-
laris (Schust.) Schust. & Damsh.; L. alpestris
auct. non (Schleich.) Evans. - Zhukova &
Rebristaya 1987)
AT: Belyy II, Khabeiyaha II, Tambey III, Tirvyyaha /,
Kharasavey II; NT: Matyuiyaha II. Tiutey 2, Venuieuo /,
Bovanenkovo 8, Tomboitosyo I, Neromayaha IV, Marre
Sale /, Yuribeitoyaha V, Khakhayayaha II, Lyakkotosyo
I; ST: Khutyyaha I, Khevesyo III, Laptayaha II, Er'yaha
II,
Khadytayaha /.
In herb-willow-moss, moss-lichen and dive-
rse swampy tundras, herb willow stands, on
brook and lake banks. While L. major prefers
comparatively rich soils, it is often associated
with acidophilous species. Always with
gemmae; male and female plants very rare.
In Yamal the species is very malleable in
its sinus, which varies from usually crescentic
to angulate with concave or convex sides;
lobe form, which is often horn-like but
sometimes - obtuse- or acute-angled; position
of maximal width of the leaf (mostly in the
lower third, occasionally in the middle); leaf
symmetry (usually the leaves are asymmetric
- with strongly convex ventral and stright
dorsal margin, but some, mainly small forms
of the species have more or less symmetric
leaves); cell size, which ranges for median
cells from 17 - 20 x 23 - 25 mkm to 28 -
35 x 30 - 42 mkm, etc. There is a certain
overlapping of the variability ranges of
Lophozia major with L. excisa, L. sudetica
and L. rubrigemma (for differentiation see
characteristics of these species).
Lophozia cf. rubrigemma Schust. (Fig.4:1-4)
AT: Tambey I; NT: Neromayaha I, Marre Sale 1.
In Cassiope tundra on bare soil with Anthelia
juratzkana, Anastrophyllum minutum and
Blepharostoma; in willow stand on bare soil
among crustose lichens, individual stem with
Lophozia major and Scapania hyperborea
mod. parvifolia-gemmipara; in willow-moss
tundra among with Tritomaria quinqueden-
tata, Ptilidium ciliare, Scapania scandica.
With gemmae. The scant Yamal material of
the species and rather poor our knowledge of
it on the whole as well as the overlapping of
variability ranges with Lophozia major cause
serious difficulties with identification of L,
rubrigemma and some problems remain un-
soluble.
The plants from Tambey share with typical
L rubrigemma leaf form and leaf position on
the stem but differ in comparatively small
cells and gemmae (Potemkin 1990a). The
plants from Neromayaha are very similar to
the type (RMS 45961c), which was investi-
gated for comparison. They have the same
leaf form, large cells (marginal 23 -
38 x 20 - 34 mkm, median 28 -
37(45) x 23 - 31 mkm) and gemmae, 23 -
31 x 17 - 28 mkm to 34 - 41 x 2028 mkm,
but material is very sparing, The last speci-
men, from Marre Sale, is also problematic.
The plants are large-ceiled with pellucid
leaves, resembling L. excisa more rather than
L.
rubrigemma Cells: however, often develop

The Hepaticae of the Yamal Peninsula 75
100 jim
Fig. 4. Lophozia cf. rubrigemma (1
Cells of leaf margin; 4 - Leaves; 5 -
Scale bars: 1 mm - for 1, 4; 0.5 mm
Khevesyo (After Potemkin 1990a).
- 4) and
Cryptocolea
imbricata (5, 6). 1 - Shoot, lateral aspect; 2 - Gemmae; 3 -
Longisection through gynoecium; 6 - Margin of inner bract with slime papillae,
for 5; 100 mkm - for 6; 50 mkm - for 2, 3. 1 - 4 - from Tambey; 5 - 6 - from
somewhat bulging trigones as in L.
rubrigemma, which do not occur in L. excisa.
Lophozia pellucida Schust. var. minor
Schust.
NT:
Neromayaha II, Khakhayayaha I; ST: Khutyyaha
I. It is a true basiphilous species of "difficult
sites",
growing usually on nival clay and
loamy slopes on bare soil with Nardia
geoscyphus, Barhilophozia quadriloba,
Cephatozia pleniceps, Tritomaria quinque-
dentata, Lophozia excisa, L. major, L. hetero-
colpos var. arctica, Jungermannia polaris, J.
obovata s.l, etc.
Lophozia pellucida var. minor is repre-
sented in Yamal mostly by very small forms
'mod. angusifolia parvifolia-fusco vel viridis),
which are often only 0.3 - 0.4 mm wide and
1-3 mm long. They have subisodiametric to
rectangular, not strongly elongated cortical
cells,
14 - 25 x 25 - 70 mkm, and small leaf
cells,
often 17-23 mkm only. Such forms
are easily distinguished due to large, 25 -
36(42) x 20 - 31 mkm, polygonal, angulate,
colourless to yellow and brownish, gemmae,
exceptionally with the traces of violet pig-
mentation; thin-walled cells almost without
trigones; stem about 10 cells high with very
weak dorsiventral differentiation of the
medulla, often blackish ventrally; violetish
fuscous bases of rhizoids. Such small forms
of the species may be easily missed when
they grow in small quantity among other
hepatics.

76 A. D. POTEMKIN
1 mm
100 дт
Fig. 5. Gymnocolea
fascinifera.
1 - Leaves; 2 - Sector of shoot of mod. angustifolia, antical aspect; 3 - Sector of
shoot of mod. latifolia, poetical aspect; 4 - Male bract; 5 - Cross section of stem with unusually thick-walled cortical
cells; 6 - Underleaf on stem; 7 - Median cells. Scale bars: 1 mm - for 1 - 4; 100 mkm - for 5 - 7. 1-3, 5-7 - from
Kharasavey; 4 - from Belyy.
GYMNOCOLEA (Dum.) Dum.
Gymnocolea fascinifera Potemk. sp. nov.
(Cladopodiella fluitans auct. non (Nees)
Buch p. p. - Zhukova & Rebristaya 1986)
(Figs.
5, 6).
Species Gymnocoleae boreali (Frisv. &
Moen) Schust. pigmentationis secundariae
magnitudine ac charactere similis est, a qua
rhizoidibus fasciculatis a basibus ventralibus
foliorum et partibus caulis adjantibus abeun-
tibus, foliis manifeste inaequilobatis inte-
gerrimis, papillis mucosis dentibusque
marginalibus destitutis, haud inunctis, guttis
oleosis numerosioribus minoribus, plerumque
4-6 mkm in diam, (2)5 - 12(16) in quavis
cellula locatis, caule pro more tenuiore
(cellulis (5)6 -8(11) in diam.), cellulis cor-
ticalibus leptodermaticis latioribus (20)23 -
28 (30) x (45)55 - 110(145) mkm ad
(28)30 - 34(38) x 30 - 40 mkm in parte
caulis dorsali sitis differt.
Plants green to brown, not glistening, 1 -
2.5 mm wide and 5-15 mm long, often fur-
cate terminal and rarely ventral and lateral
intercalary branched. Stem (5)6 - 8(11) cells
high; cortical cells weakly differentiated from
cells of medulla, rather thin-walled, usually
strongly elongated, dorsally (20)23 -
28(30) x (45)55 - 110(145) mkm, but spo-
radically subisodiametric and then wider,
(28)30 - 34(38) x 30 - 40 mkm. Rhizoids
sparse, in more or less distinct fascicles from
ventral leaf base and adjacent part of stem
(leaves detach with rhizoids). Leaves usually
asymmetric with shorter and narrower antical

The Hepaticae of the Yamal Peninsula 77
Fig. 6. Gymnocolea fascinifera. 1-4 - Leaves; 5 - Cross section of stem; 6 - Median cells with oil-bodies. Scale bars:
0.5 mm - for 1 - 4; 100 mkm - for 5; 20 mkm - for 6. All from the holotype.
lobe,
widest in the middle third, more often
a little longer than broad, but in general very
malleable in respect of width/length ratio,
which is ca. 1:0.7 - 2.0; sinus wide, V- or U-
shaped, ca.
1/3-1/2
the leaf length; lobes
from acute to apiculate, with 1 - celled
apiculus formed by subisodiametric cell, to
obtuse and rotund; margins entire, occasion-
ally more or less crooked, without any teeth
or slime papillae. Amphigastria small,
onlyoccasionally discernible, mainly when
purple pigmentation developed, formed of two
stalked slime papillae. Cells more or less
thin-walled, with small concave-sided to con-
spicuously bulging trigones, variable in size:
marginal (tangentially measured in distal
half of leaf) 17-31 mkm, median and basal
18 - 36 x 23 - 45(55 - 70) mkm. Oil-bodies
small, granulate, spherical, 4 - 6(7.5) mkm
in diam., exceptional individuals oval, up to
7-9x5-6 mkm, (2)5 - 12(16) per cell.
Cuticle smooth to usually striolate.
Dioicous. Androecia long, spicate. Male
bracts strongly concave, very asymmetric,
with small antical lobe, 1 - 2 - androus. Fe-
male plants unknown.
Holotype: USA, Alaska, Seward Peninsula,
south Killeak Lake, in well developed troughs
of high centered polygon, with Scapania
paludicola, July 28, 1992 A. D. Potemkin,
coll. № 92-97 01 (LE).
Besides the type locality where the species
was collected at first in fresh state, with oil-
bodies, it is known from one more locality in
Alaska, north of the Seward Peninsula,
neighbourhood of Kotzebue (with androecia)
(Potemkin 92 - 132 03) and from the fol-
lowing places in Yamal:
AT:
Belyy I, Kharasavey II; NT: Matyuiyaha I; ST:
Khutyyaha I, Er'yaha I.
In Sphagnum tussock bog and bogs with
flowing water, in herb-willow, grass-cotton
grass and sedge-lichen-moss tundras, often
among Drepanocladus and Sphagnum, with
Scapania paludicola var. rotundiloba,
Ptilid-
ium ciliare, Pseudolepicolea fryei, Barbi-
lophozia kunzeana, B. binsteadii, Gymnocolea
inflata, Odontoschisma elongatum, Ble-

78 A. D. POTEMKIN
pharostoma, etc. Once with androecia
(Belyy).
Gymnocolea fascinifera may be confused
with the other species of the genus and with
Cladopodiella fluitans. It differs from all of
them in rhizoid position in the ventral leaf
base area - a unique feature, not found in
the other northern species of Lophozioideae
(even if only few rhizoids present at least
some of them detach with the leaf base). The
rhizoids, detaching with the leaf base, are
absolutely noncharacteristic for G. inflata
and С fluitans and only individual rhizoids
occasionally may be detached with the leaf
base of G. borealis. Besides this character G.
fascinifera differs from G. inflata in its leaf
cells with more numerous oil-bodies, mostly
yellowish brown coloration of pigmented
forms (never fuscous black), leaves usually
with distinctly unequal, sometimes apiculate
lobes and the often larger size of plants;
from G. borealis - in leaves with distinctly
unequal lobes and entire margins without
slime papillae and teeth, not glistening ap-
pearance, numerous oil-bodies ((2)5 - 12(16)
vs 1 - 6(8) per cell), often thinner stem and
thin-walled wider cortical cells, and ecologi-
cal behaviour. The principal feature, differing
G. fascinifera as well as the other species of
the genus from Cladopodiella fluitans is the
presence of terminal furcate branching.
Moreover the species under consideration has
usually smaller, collenchymatous cells and
only sporadic small amphigastria,
Gymnocolea inflata (Huds.) Dum. var. in-
flata (G. inflata var. heterostipa (Carr. &
Spruce) K. Muell., Cladopodiella fluitans
auct. non (Nees) Buch p.p. - Zhukova & Re-
bristaya 1986; Andrejeva 1981)
AT:
Belyy H, Tambey I, Kharasavey I; NT:
Matyuiyaha II, Bovanenkovo 3, Tombottoeyo HI, Mantyto
Ш, Yuribeitoyaha I, Khakhayayaha I; ST: Khutyyaha
IV, Khevesyo III, Laptayaha I, Kharangyneto 4, Er'yaha
II,
Khadytayaha 6.
In sedge, cotton grass and sedge-moss
bogs,
sedge-lichen, cotton grass and diverse
moss tundras, late snow areas. Plants with
caducous perianth and androecia rather fre-
quent.
The speceis is malleable. On bare exposed
soil it develops mod. latifolia-densifolia-сol-
orata, or more rarely - mod. parvifolia-laxi-
folia-colorata and mod. angustifolia-laxifolia-
colorata; growing in moss tufts - mod. laxi-
folia-angustifolia-viridis and mod. parvifolia-
laxifolia-viridis.
Gymnocolea inflata var. acutiloba (Kaal.)
S. Arn.
NT:
Tofflboitosyo I, Sebayaha I; ST: Khutyyaha II.
Usually on thinly turfed ground or on peat
in pools of diverse, mostly sedge and cotton
grass bogs and wet tundras, once in sedge-
lichen nival tundra, associated with
Cephalozia bicuspidata, Lophozia ventricosa
s.l., Nardia insecta, Odontoschisma elonga-
tum,
Scapania hyperborea, S. tundrae, Gym-
nomitrion apiculatum, etc. Sporadically with
sterile perianths, once with traces of
gemmae.
After investigation of Yamal and South
Greenland material (Schuster & Damsholt
82 - 1047, 82 - 1118, 82 - 1203, 82 - 1505,
82 - 1817, 82 - 2016, etc.) I have accepted
Schuster's and Damsholt's interpretation of
"Gymnocolea acutiloba" (Schuster 1988) but
treat it here as a variety because we have not
enough data on its distribution to say about
subspecies. Their interpretation gave a set of
new characters of this taxon in the Arctic
(distinctive as well as shared with var. in-
flata):
often purplish pigmentation; caducous
perianth, pluristratose in basal half with
longer teeth of the mouth, 1 - 2(3) cells
long; leaves usually widest below the middle;
leaf bases sporadically with tooth; ca. 5% of
cells with 1-2 large oil-bodies, 50-70% of
cells with 3-7 oil-bodies. Moreover G.
inflata var. acutiloba may develop rather
thick-walled leaf cells with distinct,
sometimes bulging trigones and reproduce via
gemmae. Unfortunately most of these criteria
are not constant, occur in diverse
combinations and often it is not easy to
separate var. acutiloba from var. inflata
because of the overlapping of their variability
ranges. Nevertheless typical G. inflata var.
acutiloba may grow on bare soil and in
helophytic condition, among Sphagnum for
instance (Schuster 1988), that means it is a
distinct genetic entity.
It is necessary to stress that the Arctic

The Hepaticae of the Yamal Peninsula 79
phases of var. acutiloba are more plastic in
cell size, which in the middle of the leaf is
(17)20-28x23-31(35) mkm usually, and,
consequently, often have cells larger than the
plants described from temperate regions.
Somewhat tentatively I assign to var. acu-
tUoba the plants from Sebayaha. They share
with this taxon size, leaf shape, character of
pigmentation, smooth cuticle, but differ from
it in larger cells (median leaf cells 28 - 35
mkm wide, cortical - 21 - 30 mkm wide) and
lustrous appearance, sporadic stalked slime
papillae near leaf and particulary bract base
(in typical plants - teeth), longer teeth of pe-
rianth mouth composed of 2-4
superimposed cells. I think it may be a
polyploid derivative of var. acutiloba. For
clarification of the true taxonomic position of
these plants an additional material is
necessary.
Because of the presence of slime papillae
near the leaf base, the long teeth of the
perianth mouth, and large cells and lustrous
appearance, these plants may be confused
with Gymnocolea borealis. They differ from
this species in fuscous black pigmentation
often with traces of purple pigment, smaller
size,
smooth cuticle, caducous perianth and
ecological behaviour.
TRITOMARIA Schiffn. ex Loeske
For characterises and differentiation of the
taxa of Tritomaria in Yamal, grounds for re-
jection of sectio Tritomaria and sectio
Trilophozia (Schust.) Schust. and T.
exsectiformis subsp. arctica Schust. as well as
for description of sporophytes of T.
heterophylla and androecia of T.
quinquedentata f. gracilis see Potemkin
(1990d),
Tritomaria exsectiformis (Breidl.) Schiffn.
ex Loeske s.l. (incl. subsp. arctica Schust.)
NT: Matyuiyaha I, Tomboitoeyo I, Neromayaha I,
Lyakkotosyo II; ST: Khevesyo I, Er'yaha I.
In diverse moss and moss-lichen tundras,
individual shoots and tufts among mosses, as
an exception on bare soil among lichens, with
Barbilophozia binsteadii, Anastrophyllum
minutum, Lophozia ventricosa s.l., Ptilidium
ciliare, Tritomaria quinquedentata and Ble-
pharostoma. Always with gemmae
Tritomaria scitula (Tayl.) Joerg.
AT: Belyy II (Zhukova & Rebrlstaya 1987),
Khabeiyaha I: NT: Neromayaha I, Yuribeltoyaha Ш.
In willow-Dryas tundras and horse-tail
willow stands with Lophozia heterocolpos,
Blepharostoma, Scapania cuspiduligera, S.
gymnostomophila, Barbilophozia quadriloba,
Tritomaria quinquedentata, Arnellia fennica,
etc.
Always with gemmae.
Tritomaria heterophylla Schust. f. hetero-
phylla
NT: Khakhayayaha I; ST: Khevesyo Ш, Laptayaha Ш.
Usually on spots in wet moss spotty tun-
dras with Blepharostoma, Nardia geoscyphus,
Cephalozia pleniceps, Barbilophozia
quadriloba, Arnellia fennica, Lophozia
grandiretis, Jungermannia polaris, J. obovata,
Anastrophyllum minutum, Tritomaria quin-
quedentata, etc. Almost always with gemmae;
male and perianth bearing plants rather fre-
quent; seen twice with mature capsules.
Potemkin (1990d, fig. 1) provides the illus-
tration of this form by material from Yamal.
Tritomaria heterophylla f. anomala Potemk.
(Bot. Zhurn. 75(12): 1746, 1990)
' ST: Khevesyo П.
On spots in sedge-lichen dwarf shrub-moss,
grass-moss and lichen-moss tundras. The set
of associated species is similar to that of type
form. Once f. heterophylla and f. anomala
were found near each other, but in somewhat
different microhabitats. With gemmae, peri-
anths and androecia.
The most characteristic features of this
form are fuscous black secondary pigmenta-
tion; often obtuse to rotund leaf lobes; non
gibbous sinuses; comparatively small
trigones; female bracts with entire margin
and obtuse to shortly apiculate lobes; suben-
tire perianth mouth with obtuse apices of
marginal cells; fuscous to violet fuscous,
often strongly angulate and then larger
gemmae, (20)25 - 40(48) x 17 - 40 mkm.
Potemkin (1990d, fig. 1) provides the
illustration of this form by material from
Yamal.
Tritomaria quinquedentata (Huds.) Buch
Damsholt (1982) and Schuster (1988) dis-
tinguish, on the base of perianth characters,
two Holarctic subspecies of Tritomaria quin-

80 A. D. POTEMKIN
quedentata - subsp. quinquedentata and
subsp. turgida (Lindb.) Damsh. Analysis of
Siberian material of the species shows, how-
ever, that proposed distinctive characters, i.e.
perianth mouth ciliate vs shortly dentate; pe-
rianth unistratose and plicate on the most of
its length vs bistratose at least to the middle
and weakly plicate in upper part; purplish
pigmentation absent vs present; cells small vs
large, etc. occur in diverse combinations and
do not provide a foundation for separating
"subsp." turgida from "subsp." quinqueden-
tata (see for instance the description of the
perianth of T. quinquedentata var. quinque-
dentata f. gracilis below). Therefore, follow-
ing Schljakov (1980) I do not distinguish var.
or subsp. turgida and include it in var. quin-
quedentata.
Tritomaria quinquedentata var. quinque-
dentata f. quinquedentata (T. heterophylla
auct. non Schust. - Zhukova & Rebristaya
1986;
Gribova & Potemkin 1988)
AT
V; NT IV - V; ST: IV - V.
In wet, mainly moss, tundras, moss bogs,
occasionally in late snow areas. Male and pe-
rianth bearing plants sporadic; seen once
with mature capsules.
Tritomaria quinquedentata var. quinque-
dentata f. gracilis Schust.
AT:
Belyy I, Khabeiyaha II, Tambey II; NT: Tlutey /,
Bovanenkcm)
I, Neromayaha IV, Mantyto I, Marre Sale
/, Saleteyaha S, Yuribeitoyaha II, Khakhayayaha II,
Lyakkotosyo
II; ST: Khutyyaha I, Khevesyo II, Lap-
tayaha
I.
On steep herb slopes, in spotty tundras, oc-
casionally in moss bogs. Perianths and an-
droecia very rare. Androecia from shortly to
longly spicate, composed of 3 to 12 pairs of
1 - 2 - androus bracts; antheridia on 1 -
seriate stalk (17)20 mkm in diam. Only one
plant with perianth has been seen. Female
bracts similar to sterile leaves but a little
broader than long. Bracteole equal to bracts
in length, Ungulate, apiculate, coherent with
them at base. Perianth pluriplicate, 2 - 3 -
stratose at base, unistratose on the most of
its length; its mouth tabulate, crenulate with
solitary 1 - 2(3) - celled teeth
("var.
turgida"
type!).
This form of the species is most character-
istic for "difficult sites". Smallest phases of it
may develop the individual bilobed leaves,
which are not typical for the species.
Potemkin (1990d, fig. 2) provides the illus-
tration of this form by material from Yamal.
Tritomaria quinquedentata var. grandi-
gemma Potemk.
[T. quinquedentata var. grandigemma
Potemk. Bot. Zhurn. 75(12):1748, 1990,
nom. invalid, (sine holotypo). Holotypus:
Paeninsula Jamal, regio ripae sinistrae fl.
Laptajaha adjacens (68° 20' lat. bor., 73" 15'
long, orient.) in tundra Nanobetuloso-
caricosa muscosa, paludosa tumulis et
demissionibus tecta, 23.VII.1979, O.V.
Rebristaja (LE)] (Tritomaria quinquedentata
auct. non (Huds.) Buch - Zhukova &
Rebristaya 1986).
AT:
Kharasavey I; NT: Matyuiyaha I, Tiutey 1, Bova-
nenkovo
/, Sebayaha /, Khakhayayaha II; ST:
Khutyyaha
II, Khevesyo II, Laptayaha II, Er'yaha I.
In diverse moss, moss-lichen and dwarf
shrab tundras. Often with gemmae; perianths
and androecia rather rare.
The taxonomic status of var. grandigemma
remains somewhat problematic. This variety
is characterized by frequent production of
comparatively large, ca. (17)22 - 29(31 -
34) x 16 - 20(25) mkm, 1 - 2(3 - 4) - celled
gemmae and leaves with strongly gibbous si-
nuses.
On the other hand the plants are usu-
ally subpellucid with cell size and oil-body
number a little larger than in the type
variety sensu stricto and somewhat resemble
in this respect "var. turgida", the gemmae of
which have never been described. I have,
however, also never seen the gemmae of var.
quinquedentata and, consequently, certain
doubts arise about the constancy of their
size,
15 -20 mkm, which listed in all known
literature.
This taxon as well as the type variety is
polymorphous and varies from mod. grandi-
folia-laxifolia-viridis to mod. parvifolia-sub-
angustifolia-subaequiloba-colorata. Plants of
mod. parvifolia-densifolia-subaequiloba-color-
ata-gemmipara resemble very much Trito-
maria heterophylla. They differ from this
species in gemmae with usually round,
weakly projecting angles (if the gemmae have
acute, strongly projecting angles then cell
walls in angles are weakly thickened);

The Hepaticae of the
Yarmal
Peninsula 81
smaller cells in leaf lobes (excepting the
leaves with reduced lobes, resulting from in-
tensive gemma production); somewhat less
symmetric leaves; different type of the peri-
anth mouth (ciliate-dentate vs crenulate-
dentate). Potemkin (1990d, fig. 2) provides
the illustration of this variety by material
from Yamal.
ANASTROPHYLLUM (Spruce) Steph.
AnastrophyUum saxicota (Schrad.) Schust.
AT:
Tambey П; NT: Matyuiyaha I, Mantyto I; ST:
Khevesyo
I, Kharangyneto 2.
In sedge-lichen, often wet tundras and in
frost clefts and depressions of polygonal-frost
cleft tundra, associated with Barbilophozia
binsteadii, B. kunzeana, Ptilidium ciliare,
Tetralophozia setiformis, etc. See also
Potemkin (1990a).
Anastrophyllum sphenoloboides Schust.
NT:
Matyuiyaha I, Neromayaha I, Mantyto I,
Lyakko-
tosyo
I;
ST: Khutyyaha I, Laptayaha I.
In wet moss and dwarf shrub-lichen nival
tundras, on steep slope; growing in moss
tufts associated with Anastrophyllum minu-
tum,
Barbilophozia binsteadii, Lophozia ven-
tricosa s.l., Ptliidium ciliare, etc., but on
thinly turfed soil - with Cephalozia bicuspi-
data, Pleurocladula albescens, Anthelia ju-
ratzkana, Calycularia laxa and Gymnocolea
inflata. With gemmae, perianths and mature
capsules.
The species rather often occurs on thinly
turfed soil as mod. parvifolia-colarata-gem-
mipara. The plants of this modification are
very small, 0.2 - 0.4 mm wide and 2 - 4 mm
long, with thinner stem, 6-7 cells high;
scattered and short teeth of perianth mouth,
1
(2) cells long; antheridia on 1 - or 2 -
seriate stalk; gemmae 1(2)-celled, 14-20
(23) x 12 - 17 mkm, from round to 3 - 4 -
angulate, mostly with weakly projected an-
gles,
from faintly to deep purple; gemmi-
parous leaves usually strongly modified,
dentate, as in Lophozia bicrenata (see
Potemkin 1993b). Deviations from the de-
scription of the species by its author
(Schuster 1969; Schuster & Damsholt 1974)
that are mentioned above result mainly from
the unfavorable conditions for growth on bare
soil. Predominantly 1 - celled gemmae and
modified gemmiparous leaves result ap-
parently from very abundant gemma produc-
tion.
Anastrophyllum sphenoloboides may be
confused with some forms of A minutum de-
veloping purplish pigmentation. Such forms,
however, have features typical of A minu-
tum:
wide leaves, usually not longer than
broad, with wide sinus; small, not prominent
trigones and marginal cells considerably
smaller than median.
AnastrophyUum minutum (Schreb.) Schust.
var. minutum f. minutum
AT
V; NT V; ST V.
The most common Yamal hepatic, occurring
in diverse moss tundras, where it is more
frequent among Dicranum, rarer in lichen
and spotty tundras, and exceptionally on
steep slopes. Sporadically with gemmae, peri-
anth and androecia.
A polymorphous taxon. The most paradoxi-
cal deviations from phenotypes described
earlier are phases from dry sites with Anas-
trophyllum michauxii-like squarrose antical
lobes,
that resemble very much A japonicum
Steph. phenotypes of A michauxii. There are
numerous transitional forms between these
phases and typical A minutum, however.
The smallest phenotypes were found on ex-
posed bare soil. They are only 0.25 mm wide
and 2-3 mm long. Plants of mod. parvifolia-
sublaxifolia-colorata somewhat resemble
Cephaloziella arctica, of mod. parvifolia-den-
sifolia-colorata - Gymnomitrum.
Anastrophyllum minutum var. minutum f.
cuspidata (Kaal.) Schust.
AT:
Belyy I; NT: Matyuiyaha П, Bovanenkovo 1,
Neromayaha
III, Mantyto I, Marre Sale /, Yuribeitoyaha
I,
Khakhayayaha I, Lyakkotosyo П; ST: Khutyyaha Ш.
In wet and shade sites among mosses and
lichens in diverse communities characteristic
for the type form.
AnastrophyUum minutum var. grandis
(Lindb.) Schust.
AT:
Khabeiyaha I, Tambey II; NT: Matyuiyaha I,
Tomboitosyo
П, Neromayaha П, Mantyto I, Marre Sale
II, Ymibeltoyaha Ш, Khakhayayaha II; ST: Khevesyo
II,
Laptayaha II, Kharangyneto 2, Er'yaha II.
In diverse tundras characteristic for the
type variety, occasionally together with it.
Schljakov (1980) noted that it is a form of

82 A. D. POTEMKIN
wet sites with comparatively poor nour-
ishment. In Yamal, however, var. grandis is
often associated with basiphilous taxa, For
instance, in Yuribeitoyaha it was collected
with Scapania cuspiduligera, S. praetervisa,
Tritomaria scitula, Lophozia heterocolpos, L.
major, Odontoschisma macounii, etc. Once
(Laptayaha) with gemmae. Gemmae 3 -
angled to polygonal, brownish red and
colourless mixed, 16 - 20(23) x 14 - 17(20)
mkm.
This variety may be easily confused with
Anastrophyllum cavifolium. It differs from
this species in its small marginal leaf cells,
which usually considerably smaller than me-
dian and often form a discrete border; more
narrow cortical cells; rather regular, in cres-
centic rows, cell position in leaf lobes; not
distinct, smaller trigones; fewer oil-bodies,
etc.
Unfortunately the separation of both
taxa is complicated by occurrence of transi-
tional forms with features of both taxa (cf.
Zinovjeva 1969). So var. grandis may develop
very concave and individual 3 - lobed leaves
as well as very thick stems, to 14 cells high,
etc.
Anastrophyllum cavifolium (Buch & S.
Arn.) Lammes
AT: Kharasvey I; NT: Neromayaha I, Mantyto I.
In moss (Dicranum) tundras, on tussock in
yemik-sedge tussock bog with PtUidium cil-
iare, Anastrophyllum minutum, Tritomaria
quinquedentata, Barbilophozia kunzeana, Ble-
pharostoma, etc.
Subfam. MESOPTYCHIOIDEAE Schust.
MESOPTYCHIA (Iindb.) Evans
Mesoptychia sahlbergii (Iindb. & H. Arn.)
Evans
ST: Khutyyaha I.
In moss willow stand among Drepanocladus
sp.
and Paludella squarrosa with Lophozia
gillmanii, L. heterocolpos var. arctica, Pla-
giochila
porelloides, Aneura pinguis, etc.
Subfam. JUNGERMANNIOWEAE
JUNGERMANNIA L.
Subgen. JUNGERMANNIA
Jungermannia pumila With.
ST: Khevesyo I.
In moist dwarf shrub-sedge-moss tundra on
bare soil with Cephalozia bicuspidata and
Nardia geoscyphus.
Jungermannia polaris Lindb.
AT: Belyy I, Khabelyaha I, Tambey II, Kharasavey I;
NT: Tiutey J, Yuribeitoyaha I, Khakhayayaha I; ST:
Khevesyo I, Laptayaha II, Er'yaha I.
On spots in spotty tundras, rarely on bare
soil on herb and nival slopes. Often with pe-
rianths, occasionally with mature capsules.
Subgen. SOLENOSTOMA (Mitt.) Amak.
Jungermannia confertissima Nees
AT: Tambey I; NT: Tomboitosyo I, Neromayaha I,
Khakhayayaha 1; ST: Khevesyo II, Laptayaha II,
Er'yaha I.
On bare and thinly turfed soil in spotty
lichen-moss tundras, wet herb-moss willow
stands, on nival slope; associated with Nar-
dia geoscyphus, Cephalozia bicuspidata, C.
pleniceps, Aneura pinguis, Blepharostoma,
Lophozia grandiretis, Odontoschisma ma-
counii, Tritomaria heterophylla, Cryptocolea
imbricata, Anthelia juratzkana, etc. Often
with perianths, once with mature capsules.
Jungermannia jenseniana Grolle
NT: Khakhayayaha I; ST: Khevesyo II.
On spots of bare sandy ground in spotty
dwarf shrub tundras with Anthelia ju-
ratzkana, Marsupella sprucei, Prasanthus
suecicus, Cephalozia bicuspidata, Nardia
geoscyphus, etc. Twice (Khevesyo) with peri-
anths.
Yamal material of
Jungermannia
jenseniana
is too sparse to form a true notion on this
species. Phases without distinct red pigmen-
tation are difficult to separate from /.
sphaerocarpa var. nana also developing a pe-
rigynium. Differentiation from /. confertis-
sima is more problematic. The main distinc-
tive feature of the last species is rhizoid po-
sition. This character, however, is not very
constant - rhizoids may migrate to ventral
leaf base and perigynium in some forms of /.
sphaerocarpa var. nana (Schuster &
Damsholt 1974) and, moreover, in J. jense-
niana itself - in an isotype of the species
from JE (V. Schiffner. Hepaticae eur. exs.
1380.
Aplozia pusilla CJens. Orig. Ex.!)
leaves often detach with rhizoids. As a con-
sequence, the problem of taxonomic status of
/. jenseniana is critical.

The Hepaticae of the Yamal Peninsula 83
Jungermannia sphaerocarpa Hook.
var.
sphaerocarpa
AT:
Beiyy 1; ST: Khevesyo III, Laptayaha I.
On bare ground of creek, river and lake
banks with Pleurocladula albescens,
Cephalozia bicuspidata, Nardia geoscyphus,
Lophozia cf. savicziae, L. opacifolia, Anthelia
juratzkana, Blasia pusilla, etc. Almost always
with perianths, once with mature capsules.
Jungermannia sphaerocarpa var. nana
(Nees) Frye & Clark (Solenostoma pumilum
subsp. polaris auct. non (Lindb.) Schust. -
Zhukova & Rebristaya 1986)
AT:
Belyy K, Tambey П, Kharasavey П; NT:
Matyuiyaha
II, Neromayaha I, Yuribeitoyaha I,
Khakhayayaha
II; ST: Khevesyo IV, Laptayaha I.
On spots of bare ground in wet spotty tun-
dras,
rarer at the foot of steep herb and nival
slopes and on banks of creeks and lakes. Al-
most always with perianths, often with ma-
ture capsules.
Small forms of Jungermannia sphaerocarpa
var. nana, are sometimes confused with /.
polaris. They differ from the last species in
their larger cells, especially marginal, which
near leaf apex are at least 20 - 23(25) mkm
(vs 15-17, rarely 21 mkm long in /. po-
laris);
subrotund, plane or slightly concave,
or, occasionally, weakly deflexed backward
leaves, briefly decurrent dorsally and ven-
traily, oriented in parallel to stem and, con-
sequently, jutted out of the ventral surface of
the stem (in /. polaris leaves mostly ovate,
widest at base, nondecurrent, concave, ori-
ented distinctly at the angle to stem and
never jutted out of the ventral surface of the
stem).
Jungermannia sphaerocarpa var. nana in
Yamal often develops a distinct low perigy-
nium and the gametophyte lacks red or pur-
plish pigmentation while capsules and spores
are usually purplish or purplish brown.
The taxonomic status of Jungermannia
sphaerocarpa var. nana is rather problematic.
Quite possibly it is mainiy an ecologically
determined form of tundra and alpine
regions (cf. Schuster 1969; Koponen et al.
1977).
Subgen. PLECTOCOIMA (Mitt.) Amak.
Jungermannia hyalina Lyell (Solenostoma
pumilum auct. non (With.) K. Muell. -
Zhukova & Rebristaya 1986)
AT:
Khabeiyaha I, Kharasavey I; NT: Matyuiyaha П,
Saletayaha
/, Khakhayayaha II; ST: Khutyyaha H,
Khevesyo
II, Er'yaha I.
In herb-moss, herb-horse-tail and sedge
willow stands, at the foot of steep herb slope
at margin of swampy sedge depression, on
sandy bank, on degraded Sphagnum. An-
droecia, perianthas and mature capsules rare.
Reports of the species from the arctic tundra
are northernmost known.
In Yamal Jungermannia hyalina occurs as
mod. laxifolia-leptoderma-viridis, more rarely
- mod. laxifolia-mesoderma- subcolorata vel
viridis. The plants have pellucid, often
strongly asymmetric leaves, longly decurrent
dorsally; mostly colourless rhizoids, not
rarely in distinct fascicles from ventral leaf
bases;
thin-walled medulla of stem; and spo-
radically rather low perigynium. While these
characters are usually not mentioned for the
species, they often occur in /. hyalina
throughout its range (as was found in the
course of investigation of numerous collec-
tions of the species in LE).
Jungermannia obovata Nees s.l.
(Jungermannia subelliptica (Lindb. ex Kaal.)
Lev.)
AT:
Khabeiyaha II, Kharasavey II; NT: Neromayaha I;
ST:
Khutyyaha I, Khevesyo III, Laptayaha II, Er'yaha I.
In dwarf shrub sedge-(cotton grass)-moss
sedge-lichen-moss and diverse spotty tun-
dras,
on wet thinly turfed loamy soil near
temporary water courses, in late snow areas,
usually on thinly turfed soil, associated with
Anthelia juraizkana, Nardia geoscyphus,
Jungermannia polaris, J. sphaerocarpa var.
nana, Cephalozia bicuspidata, C. pleniceps,
Scapania obcordata, Tritomaria heterophylla,
etc.
Almost always with perianths, often with
mature capsules.
NARDIA S.Gray
Nardia japonica Steph.
ST:
Khutyyaha I.
In dwarf shrub-cotton grass moss tundra
and on high-centered polygon, on bare loamy
sandy soil with humus, associated with
Cephalozia bicuspidata, Gymnocolea inflata

84 A. D. POTEMKIN
var. acutiloba, Calycularia laxa and Nardia
insecta. Only male plants were found.
When the species grows with Nardia insecta
it distinctly differs in smaller size (0.2 - 0.5
mm vs 0.5 - 1.6 mm wide); underleaves,
which longer than the half of leaf length vs
considerably shorter than the half of leaf
length; and sex distribution (dioicous vs
paroecious).
Nardia geoscyphus (De Not.) Lindb. var.
geoscyphus
AT: Belyy II, Khabelyaha II, Tambey II, Kharasavey
Ш; NT: Matyulyaha I, Tiutey /, Neromayaha IV, Sale-
tayaha 4, Yuribeltoyaha I, Khakhayayaha Ш, Lyakkoto-
syo I; ST: Khutyyaha Ш, Khevesyo V, Laptayaha V,
Er'yaha I.
In moist, more rarely in comparatively dry
spotty tundras, on steep herb and nival
slopes, on banks of diverse water courses and
lakes,
in bog communities; on sandy, loamy
and peaty ground. The most common species
of Jungermannioideae on the peninsula. Of-
ten with perianths and mature capsules.
Nardia geoscyphus var. bifida Schust.
NT: Neromayaha I.
On loamy sand landslip nival slope with
Anastrophyllum minutum.
Nardia insecta Lindb.
NT:
Mantyto I; ST: Khutyyaha II.
On high-centered polygon, yernik Sphag-
num hummock and polygonal spotty- hum-
mock dwarf shrub lichen tundra; on loamy
and loamy sandy soil with humus, with
Cephalozia bicuspidata, Anthelia juratzkana,
Gymnocolea inflata var. acutiloba, Junger-
mannia jenseniana, Gymnomitrion apicula-
tum,
Marsupella sprucei, Nardia japonica
and Lophozia ventricosa s.l. With perianths
and mature capsules.
Yamal phenotypes of the species are not
typical. They usually have oval, rotund to
reniform, strongly concave, entire or mostly
retuse leaves; very deep pigmentation, from
purplish brown to fuscous or purplish black.
They share with typical plants paroecia; col-
lenchymatous large cells with granulate oil-
bodies; distinct squarrose underleaves with
rhizoids in fascicles at base, frequently nar-
rowly united with leaf bases on one side of
stem; and sporophyte characteristics. Such
forms of the species may be confused with
Nardia scalaris. They differ from it in sex
distribution; granulate, not glistening cil-
bodies; underleaves mostly connate with
leaves bases on one side of stem; often
somewhat larger cells. Moreover Nardia
scalaris is principally an amphioceanic
species. Individual reports of it from conti-
nental regions are apparently based on phe-
notypes of N. insecta with entire leaves.
CRYPTOCOLEA Schust.
Cryptocoiea imbricata Schust. (Fig. 4: 5-6)
AT: Khabeiyaha I; ST: Khutyyaha I, Khevesyo II,
Laptayaha П.
On bare ground in spotty tundras and on
herb slopes, usually with Tritomaria hetero-
phylla, Nardia geoscyphus, Jungermannia
confertissima, J. obovata, Cephalozia pleni-
ceps, more rarely with Arnellia fennica,
Jungermannia sphaerocarpa var. nana, Ble-
pharostoma, Odontoschisma macounii, Trit-
omaria quinquedentata, Prasanthus suecicus,
Gymnomitrion corallioides, and Anthelia Ju-
ratzkana. Male and female plants not rare.
For details see Potemkin (1990a).
Subfam. MYUOIDEAE Grolle
MYIIA
S. Gray
Mylia anomala (Hook.) S. Gray
NT: Mantyto I; ST: Khutyyaha Ш, Khevesyo I.
In sedge-Sphagnum tussock-pool swampy
tundras, on peat outcrop and on steep herb-
dwarf shrub nival slope. Usually with Barbi-
lophozia binsteadii, B. kunzeana, Ptilidium
ciliare, Cephalozia lunulifolia, C. bicuspidata,
Scapania irrigua, Sphagnum spp., etc. Twice
with gemmae (Khutyyaha).
In Mantyto individual plants of mod. parvi-
folia-densifolia- colorata were collected on a
nival steep slope on bare soil with Pleuro-
cladula albescens, Gymnomitrion concinna-
tum,
Cephalozia bicuspidata, C. pleniceps,
Anastrophyllum minutum, and Lophozia cf.
savicziae. The plants are only 0.8 - 1.0 mm
wide and about 3 mm long. They resemble
Nardia scalaris rather than Mylia. Such un-
usual phases of the species differ from Nar-
dia scalaris in the character of cross section
of stem with regularly distinctly projecting,
larger cortical cells, 60 - 80 x 45 - 60 mkm

The Hepaticae
of the
Yamal Peninsula 85
(Schuster
1969,
Figs. 260:8, 299:8); larger
leaf cells,
48 - 56 x 28 - 46 mkm, and un-
derleaves;
and not
persistent, more numerous
oil-bodies
of
different structure.
GYMNOMITRIACEAE Klinggr.
PRASANTHUS
(Lindb.) Lindb.
After description
of
Prasanthus jamalicus
(Potemkin 1992b)
two
species
of the
genus
became known. They differ
as
follows:
KEY TO SPECIES OF PRASANTHUS
1.
Minute (width
of
sterile shoots
200 - 350
mkm),
purplish brown
to
purplish black,
unglistening, with distant leaves
on
sterile
shoots, resembling small species
of
Nardia;
stolons rare; cell walls
of
sterile leaves thin;
basal leaf cells mostly subisodiametric
(length
:
width ratio
- 1:1.5(2)),
undifferentiated from median cells;
thickenings
of
inner capsule wall layer
nodular, often with semiannular bands
in
basal parts
of
valves; elaters
3(4) -
spiral,
exceptionally
2 - and 5 -
spiral
..sectio
PSEUDONARDIA
Potemk.,
P.
jamalicus
1.
Small (width
of
sterile shoots 350-700
mkm),
yellow
to
brown, very rarely
the
ven-
tral part
of the
perigynium purplish; glisten-
ing, with imbricate, rarely subimbricate
leaves; stolons common; cell walls
of
leaves
thick; basal leaf cells mostly elongated
(length
:
width ratio
- 1:2-3),
distinctly
differentiated from subisodiametric median
cells;
thickenings
of
inner capsule wall layer
semiannular; elaters
2 -
spiral, exceptionally
3 -spiral sectio
PRASANTHUS, P.
suecicus
Prasanthus jamalicus Potemk.
[Ann. Bot.
Fennici 29:319,
1992]
NT:
Khakhayayaha
I; ST:
Khutyyaha
I.
In pure patches
on
exposed windswept
sandy soil,
in
Khutyyaha (Holotype) with
small admixture
of
Scapania
sp.,
Prasanthus
suecicus, Marsupella sprucei, AntheUa
ju-
ratzkana. With mature capsules.
Prasanthus suecicus (Gott.) Lindb.
AT:
Bclyy
II,
Khabeiyaha
III,
Tambey
II,
Kharasavey
II;
NT:
Matyuiyaha
II,
Neronayaha
III,
Saletayaha
4,
Yuribeitoyaha
I,
Khakhayayaha
III,
Lyakkotocyo
I; ST:
Khutyyaha
III,
Khevesyo
IV,
Laptayaha
HI,
Kharangy-
neto
/,
Er'yaha
П,
Khadytayaha
2.
In spotty
and
lichen tundras, steep thinly
turfed slopes,
on
exposed windswept sandy
soil. Usually fertile, often with mature cap-
sules.
MARSUPELLA Dum.
Marsupella sprucei (Limpr.)
H.
Bern.
AT:
Khabeyyaha
1,
Tambey
I; NT:
Matyuiyaha
I,
Neromayaha
II,
Saletayaha
5,
Khakhayayaha
П,
Lyakkotoeyo
I; ST:
Khutyyaha
E,
Khevesyo
III,
Lap-
tayaha
I,
Er'yaha
I,
Khadytayaha
2.
Usually individual plants
on
windswept
sandy soil
in
spotty tundras, steep slopes,
and other similar sites, associated mostly
with Prasanthus suecicus
and
Gymnomitrion
coralUoides. Almost always fertile, sporadi-
cally with mature capsules.
GYMNOMITRION
Corda
Gymnomitrion apiculatum (Schiffn.)
K.
Muell.
(G.
corallioides auct.
non
Nees
- Gri-
bova
&
Potemkin
1988)
AT:
Belyy
I,
Tambey
I; NT:
Matyuiyaha
П,
Tomboitosyo
I,
Saletayaha
1; ST:
Khutyyaha
I,
Khevesyo
I,
Laptayaha
I.
On nival north-facing sandy slopes,
in
herb-willow Gymnomitrion
and
dwarf shrub-
lichen tundras
on
sandy soil, associated with
the other regional Gymnomitriaceae, AntheUa
juratzkana, Lophozia
cf.
savicziae,
L.
opaci-
folia, Cephalozia bicuspidata, Pleurocladula
albescens,
and
Diplophyllum taxifolium. Once
(Matyuiyaha) with perianth.
The species occurs
in
Yamal
in two
phases:
Gymnomitrion corailioides-like, dorsiventrally
compressed, with very dense, often hardly
discernible leaves
and the
other phase,
re-
sembling more Gymnomitrion concinnatum,
almost
not
dorsiventrally compressed, with
more distant, distinctly discernible leaves.
As
a consequence
it
often difficult
to
identify
the
species
for
sure
in the
course
of
superficial
investigation
of
specimens under
a
dissecting
microscope, especially because
it
very rarely
develops
red
pigmentatioin
in
Yamal.
G.
apiculatum differs from
G.
concinnatum,
when sterile,
in the
shallower
and
wider,
obtuse-angled sinus; thin-walled marginal
cells,
erose
at
least
on old
leaves; smooth
cu-
ticle;
apiculate lobes, with thin-walled,
quickly erose,
not
permanent apiculi
of 2 su-
perimposed cells
(in
G. concinnatum they
are
thick-walled,
not
erose, permanent, while

86 A. D. POTEMKIN
sometimes very similar in form), etc. The
main distinctive features from G. corallioides
are the following: apiculate leaf lobes (dis-
cernible on young leaves); wide sinus;
thinner-walled leaf cells,
Gymnomitrion concinnatum (Ughtf.) Corda
AT: Belyy Ш, Khabeiyaha I, Tambey II; NT:
Matyuiyaha II, Tiutey /, Mantyto I, Khakhayayaha I;
ST: Khutyyaha II, Khevesyo II, Kharangyneto /, Er'yaha
I, Khadytayaha 3.
In spotty (often Gymnomitrion) tundras, on
steep slopes, late snow areas, etc., on sandy
and loamy ground. Male plants and plants
with mature capsules sporadic.
Gymnomitrion concinnatum differs from the
two other regional representatives of the
genus in its narrow and more deep sinus;
faintly but distinctly papillose cuticle; and
not erose while often decolorate leaf margins.
The species varies from mod. subdensifolia-
viridis to mod. densifolia-fusca.
Gymnomitrion corallioides Nees
AT: Belyy IV, Khabeiyaha IV, Tambey IV, Kharasavey
III; NT: Matyuiyaha П, Tiutey 2, Neromayaha IV, Marre
Sale 1, Saletayaha 10, Yuribeitoyaha T, Khakhayayaha
Ш, Lyakkotosyo I; ST: Khutyyaha IV, Khevesyo III,
Laptayaha II, Kharangyneto 3, Er'yaha II, Khadytayaha
3.
Usually in spotty sedge-lichen and sedge-
Racomitrium-lichen tundras, where it is a
dominant taxon on spots (in this connection
these tundras are named Gymnomitrion or
Gymnomitrion-Racomitrium tundras). Male
plants and plants with mature capsules are
sporadic. The species varies from mod. den-
sifolia-colorata (on exposed sites) to mod.
subdensifolia-viridis (in shade).
SCAPANIACEAE Migula
SCAPANIA (Dum.) Dum.
Subgen.
JENSENIA
S.Arn.
Scapania obcordata (Berggr.)
S.Arn.
AT: Belyy Ш, Khabeiyaha II, Tambey II, NT:
Matyuiyaha I, Bovanenkovo /, Neromayaha II,
Yuribeitoyaha II. Khakhayayaha II; ST: Khutyyaha II,
Khevesyo III.
In diverse moist moss tundras, near tempo-
rary water courses, on steep herb and nival
slopes, in sedge-moss bogs with flowing wa-
ter, and on peat outcrop. Usually with gem-
mae or with traces of gemmae on modified
leaves, sporadically with androecia and peri-
anths.
Scapania obcordata is an extremely
variable species, which, when growing
on exposed bare soil usually develops
mod, parvifolia-subaequiloba-colorata but in
shade and moist conditions produces
the phenocopies of S. irrigua. The latter
phenotypes differ from S. irrigua in stem
anatomy, character of the cell net at base
and near apexes of leaves (see Schuster
1974;
Schljakov 1981), peculiar modification
of gemmiparous leaves, and gemmae and oil-
bodies characteristics. Oil-bodies in S.
obcordata are often with a more or less
distinct central droplet (Schuster 1974, Fig.
346:
2).
Subgen. KAALAASIA Buch
Scapania gyrnnostomophila Kaal. f.
gymnostomophila
NT: Yuribeitoyaha I.
In cleft on spot in spotty-cleft tundra with
Lophozia heterocoipos, L. major, Barbi-
lophozia quadriloba f. glareosa, Tritomaria
quinquedentata f. gracilis. With gemmae.
Transitional forms to Scapania gymno-
stomophila f. incurva were found. It confirms
the supposition of H. Persson (1946) and
Schuster (Schuster et al. 1959) that
"Diplophyllum incurvum Bryhn & Kaal." is a
peculiar form of Scapania gymnostomophila
characteristic for exposed sites.
Scapania gymnostomophila f. incurva
(Bryhn & Kaal.) Schust.
NT: Yuribeitoyaha II.
On bare soil in willow-Dryas spotty tundra
with Arnellia fennica, Tritomaria scitula,
Lophozia heterocoipos, L. badensis, and
Barbilophopzia quadriloba f. glareosa.
Scapania gymnostomophila f. incurva
resembles very much S. calcicola (H. Arn. &
Perss.) Ingh. subsp. ligulifolia (Schust.)
Danish. & Long, differing from this taxon in
its smaller gemmae and number of oil-bodies
which are usually 1-2 per median cell.
Subgen. BUCHIELLA Schust.
Scapania zemljae S. Arn (? S. invisa
Schust.)
AT: Belyy I; NT: Tomboitosyo I, Neromayaha III,
Khakhayayaha II; ST: Khutyyaha II, Khevesyo II,
Laptayaha II.

The Hepaticae of the Yamal Peninsula 87
Usually on spots in spotty tundras,
occasionally on thinly turfed sandy soil,
associated mostly with Gymnomitrion
corallioides, Prasanthus suecicus, Cepha-
loziella arctica, Anthelia juratzkana, Lopnozia
alboviridis, L. excisa, Marsupella sprucei,
Nardia geoscyphus, etc. In Belyy the species
was collected, however, in association with
mainly basiphilous hepatics: Lopnozia
heterocolpos var. harpanthoides, Barbi-
lophozia quadriloba, Jungermannia polaris,
Cephaloziella arctica, and Odontoschisma
macounii. Always with gemmae, occasionally
with androecia, once with perianth
(Khakhayayaha). For descriptions of male
plants and characteristic of gemmae see
Potemkin (1993b).
The species is assigned here to subgenus
Buchiella rather tentatively. Sterile plants of
Scapania zemljae have much in common with
the type species of the subgenus - S.
cuspiduligera: erect and sheathing, often
widely 2(3) - stratose leaf base; ability to
develop a border of smaller thick-walled
marginal cells; colourless to red and brown,
2 - celled gemmae; not persistent oil-bodies;
decurrent ventral leaf margin, etc. (Widely
2(3) - stratose leaf bases have not been
noted previously for the species; this
character was found during investigation of
original collections of O. Ekstam from
Novaya Zemlya, including the type of the
species (Novaya Zemlya, S. Island, Sinus
Karmakulski, 0. Ekstam, 3.09. 1901) and of
Yamal materials). The structure of the
perianth (which has been seen only once) is
different from that of S. cuspiduligera,
however. The perianth is pluriplicate in its
upper part, with (3)4 deep plicae descen-
ding to the base. The mouth is bleached,
dentate, with bluntish to triangular teeth
with a uniseriate apiculus of 1 - 2(3)
subisodiametric cells; the larger teeth arise
from a base 3-4(5) cells broad. The
perianth characteristics separate S. zemljae
from S. cuspiduligera. But these differences
cannot be evaluated firmly on the basis of
only one collection. Additional collections of
plants with perianths are needed for the
elucidation of the taxonomic position of this
species.
Scapania cuspiduligera (Nees) K.Muell.
var. cuspiduligera
AT:
Khabeiyaha I; NT: Matyuiyaha I (Zhukova &
Rebristaya 1986), Maire Sale 2, Yuribeltoyaha III; ST:
Laptayaha I.
In herb-moss and horse-tail willow stands,
sedge-yernik-willow tundra, associated with
Blepharostoma, Tritomaria quinquedentata,
T. scitula, Lophozia heterocolpos, L. major,
Barbilophozia quadriloba f. glareosa, Anastro-
phyllum minutum var. grandis. Always with
gemmae, which are 2 - celled and almost
invariably yellowish brown with a deeper
pigmented septum.
Comparative abundance of the species was
mentioned only in Yuribeitoyaha where the
manifestations of its variability from common
mod. laxifolia-viridis (plants to 10 mm long
and 3 mm wide) to mod. parvifolia-(den-
sifolia)-colorata (plants about 2 mm long and
0.7 mm wide) were observed. Plants of the
last modification differ considerably from
typical forms as was noted by Schuster also
(Schuster et al. 1959). They have deep
fuscous (rarely brown) pigmentation; leaves
with unequal, occasionally shortly apiculate
lobes,
sporadically with individual obtuse
teeth formed mostly by projecting angles of
marginal cells. The border is present only
when marginal cells are bleached. Its absence
correlates with development of deep fuscous
pigmentation. Cells are considerably smaller,
median 14 - 17 mkm, marginal 10 - 17 mkm,
occasionally with rough striolate cuticle
(elongated papillae to 3 - 4 mkm wide and
10-15 mkm long). Sheathing of the stem by
leaf bases as well as long ventral decurrence
are not distinct, especially in mod. densifolia.
Such phenotypes are usually connected with
typical ones by transitional forms. They have
characteristic bleached ventral leaf bases and
modified apexes of gemmiparous leaves,
brown, 2 - celled gemmae and, according to
Schuster & al. (1959), brown, big oil-bodies,
(1)2 - 4(5 - 6) per cell.
Scapania cuspiduligera var. diplophyllopsis
Schust.
NT:
Neromayaha П.
In yernik-willow and lichen tundras, on
loamy soil with humus, associated with
Jungermannia obovata, Nardia geoscyphus,

88
A.
D. POTEMKIN
Anthelia juratzkana, Lophozia jurensis, L.
major, L. heterocolpos, Ptilidium ciliare,
Blephwostoma, Barbilophozia hyperborea, B.
quadriloba, Tritomaria guinguedentata, T.
exsectiformis, Cephaloziella rubella subsp.
arctogena. Always with gemmae, once with
androecia (for description of androecia see
Potemkin (1993b)).
Until now the range of variability and
distinctive features of this variety remain not
quite clear. A Diplophyllum taxifolium - like
ventral lobe is not a constant character.
Gemma production often leads to reduction
of the ventral lobe, resulting occasionally in
leaves with almost equal lobes. I think, the
most important criteria, differentiating this
variety from var. cuspiduligera, is the
gemmae, green until near maturity, then
becoming brown, and leaf lobes often with
few, small sharp denticulations and a 1 -
celled mucro.
Subgen. SCAPANIA
Scapania curta (Mart.) Dum. var. curta
AT:
Belyy I (Zhukova & Rebristaya
1987),
Khabeiyaha
I,
Tambey I, Kharasavey I; NT: Neromayaha П, Mantylo
I,
Yuribeitoyaba I, Khakhayayaha I; ST: Khutyyaha I,
Khevesyo
II, Laptayahs П, Er'yaha I.
In dwarf shrub-moss and spotty tundras, at
the foot of herb and nival slopes, in flat-
polygon aad sedge-moss bogs, on peat
outcrop. OcasionalSy with gemmae, once with
perianth (Er'yaha).
Scapania curta var. grandiretis Schust.
NT:
Khakhayayaha II; ST: Khutyyaha И.
In late snow areas, tussock bog, tussock
moss tundra, associated with Tritomaria
guinguedentata f. gracilis, Nardia geoscyphus,
Jungermannia confertissima, Lophozia
opacifolia, L. cf. savicziae, Anthelia
juratzkana, Cephalozia bicuspidata, Scapania
curta var. curta, etc. Usually with gemmae,
once with perianths (Khakhayayaha).
Scapania scandica (H. Arn. & Buch) Macv.
var. scandica (S. parvifolia Warnst.; S.
mucronata auct. non Buch - Potemkin 1989)
AT:
Belyy II, Khabclyaha II, Tambey III, Kharasavey
II;
NT: Matyuiyaha П, Tomboitosyo I, Neromayaha III,
Mantyto
I. Marre Sale 1, Saletayaha /, Yuribeitoyaba I,
Khakhayayaha
II; ST: Khutyyaha V, Khevesyo V, Lap-
layaha
III, Kharangyneto 2, Er'yaha П, Khadytayaha /.
In diverse tundra communities formed by
mosses, lichens, sedges, grasses and dwarf
shrubs, on spots in spotty tundras, in late
snow areas, occasionally in bogs composed of
sedge, cotton grass and mosses (mainly by
species of Drepanocladus, Hypnum, Sphag-
num).
It is the most widespread taxon of the
genus in Yamal. Often with gemmae, spora-
dically with androecia and perianths. For
details on this and the other varieties of the
species see Potemkin (1993a).
Scapania scandica var. argutedentata Buch
ST:
Khevesyo I.
At base of steep herb slope and on moist
loamy scarp with Lophozia opacifolia, L. cf.
savicziae, Blasia pusilla, Nardia geoscyphus,
Jungermannia sphaerocarpa, Anthelia jurat-
zkana, Cephalozia bicuspidata, C. pleniceps,
Pleurocladula albescens. With gemmae,
androecia, perianths and mature capsules.
Illustrations in Potemkin (1993a).
Scapania scandica var. grandiretis
(Schljak.) Schljak.
AT:
Kharasavey I; NT: Neromayaha I, Khakhayayaha
I.
On spots in spotty tundras, on margin of
lake and in nival tundra shortgrass meadow;
associated with Scapania scandica var.
scandica, S. curta, Cephalozia bicuspidata,
Lophozia ventricosa s.l., L. excisa, L.
wenzelii, L. major, Anastrophyllum minutum,
Nardia geoscyphus, etc. With gemmae and
androecia.
Scapania mucronata Buch
NT:
Khakhayayaha I.
On bare loamy sandy ground in spotty
hillock yeraik tundra with Cephaloziella
arctica, Cephalozia bicuspidata, Odonto-
schisma macounii, Anthelia juratzkana,
Scapania cf. obcordata. With gemmae. This
report is based on sparse material. The
species is represented by mod. parvifolia-
colorata-gemmipara. Nevertheless the plants
are quite typical: brownish, without any
traces of red or purplish pigment, with
undifferentiated marginal leaf cells and
rather short keel, < 0.5 the ventral lobe in
length. In spite of intensive gemma formation
the leaf margins are entire.

The Hepaticae of the Yamal Peninsula 89
Scapania praetervisa Meyl.
AT:
Beiyy I, Khabelyaha I, Tambey I, Kharasavey II;
NT:
Matyuiyaha I, Tomboitoeyo II, Neromayaha II,
Lyakkotoeyo II; ST: Khutyyaha I, Laptayaha I.
In diverse moss, dwarf willow-moss
tundras, polygonal dwarf shrub-lichen-Gym-
nomitrion and dwarf shrub-moss-lichen
tundras, high-centered polygon and in grass-
sedge tundra. Often with gemmae, once with
androecia and juvenile perianths.
Scapania irrigua (Nees) Nees var. irrigua f.
irrigua
AT:
Belyy Ш (Zhukova & Rebristaya 1987),
Khabeiyaha I, Kharasavey I; NT: Matyuiyaha II,
Bovanenkovo /, Tomboitoeyo I, Neromayaha I,
Saletayaha 1, Khakhayayaha I; ST: Khutyyaha Ш,
Khevesyo Ш, Laptayaha Ш, Khadytayaha /.
In diverse moist sedge-moss tundras, moss-
cotton grass and sedge bogs, near banks of
water courses and lakes, etc. Gemmae,
androecia and perianths rare.
Scapania irrigua var. irrigua f. rubescens
(Buch) Schust.
NT:
Mantyto I.
In sedge bog. With androecia.
Scapania irrigua var. rufescens (Loeske)
Loeske
NT:
Bovanenkovo /.
In cotton grass-sedge-hypnum bog, among
Calliergon sarmentosum.
Scapania degenii Schiffn. ex K. Muell. (5.
hyperborea auct. non Joerg. p.p. - Potemkin
1989)
NT:
Khakhayayaha II.
In willow-sedge lichen-moss and sedge-
Hylocomiutn tundras, associated with
Ptilidium ciliare, Tritomaria quinquedentata,
Blepharostoma, Cephaloziella arctica, Lopho-
zia heterocolpos, Scapania cuspiduligera, and
Anastrophyllum minutum.
Yamal plants represent mod. grandifolia-
integrifolia - coiorata -egemmipara. Potemkin
(1989) attributed them to Scapania
hyperborea because of the absence of
gemmae, occasionally dorsally arched leaves
with entire margins and usually not apiculate
lobes of which the dorsal is comparatively
large. An analysis of the Yamal specimens of
S. degenii and the specimen from Arctic
Alaska determined by W. C. Steere July 22,
1951,
which contains plants without gemmae,
with dorsally arched, entire margined leaves
as well as rather typical plants with gemmae
and dentate leaf margins, showed that leaf
teeth formation in the species correlates with
gemma formation, resulting in apiculate leaf
lobes,
of which the ventral often becomes
more narrow.
Phenotypes of Scapania degenii without
gemmae differ from S. hyperborea in the
following features: 1) oil-bodies usually
persistent in drying; 2) keel distinctly shortly
winged (a winged keel is an exception in S.
hyperborea - I have never seen it in arctic
plants of the species, while K. Mueller
(1951 - 58) mentioned it); 3) semicircular
(vs irregular) cell position in leaf lobes; 4)
the often 3 - stratose cortex (vs 1 - 2 -
stratose, only exceptionally 3 - stratose) (for
evaluation of the last character it is necessary
to take into account habitat conditions).
Scapania hyperborea Joerg. (incl. f.
subaequalis Schust.; ? S. brevicaulis auct. non
Tayl. - Zhukova & Rebristaya 1986)
AT:
Belyy I, Khabeiyaha П, Kharasavey I; NT:
Matyuiyaha I, Bovanenkovo 1, Tomboitoeyo I,
Neromayaha I, Mantyto I, Yuribeitoyaha I; ST:
Khutyyaha П, Khevesyo I.
In sedge bogs and tundras (often of nival
type),
herb nival communities, in moss
(Hylocomium, Dicranum) and moss-lichen
tundras, on sandy margin of lake; often on
sandy ground. Sporadically with gemmae,
which vary in coloration from pink, purple
and violet to fuscous, reddish fuscous and
purple blackish fuscous; once with androecia.
For characteristics of reproduction of the
species by gemmae and for consideration of
status of f. subaequalis see Potemkin
(1993b).
Scapania hyperborea is an extremely
malleable species represented in the Arctic
by numerous phenotypes which in details
were described by Schuster (1974) and
Schuster and Damsholt (1974). Most
paradoxical of them is the Scapania hel-
vetica-phenotype. The plants reported from
Yamal as S. brevicaulis Tayl. (Zhukova &
Rebristaya 1986) apparently should be
attributed to this phenotype. They are very
small, brownish pigmented with bistratose

90 A. D. POTEMKIN
keel at least in basal half of leaves and very
characteristic for the species broadly oval to
almost spheric pinkish to purple 2 - celled
gemmae.
Scapania tundrae (H. Am.) Buch (S.
pulcherrima Schust. syn. nov.)
AT:
Belyy I, Kharasavey I; NT: Matyuiyaha П,
Bovanenkovo
I, Tomboitosyo I, Neromayaha Ш, Mantyto
П,
Saletayaha /, Khakhayayaha П; ST: Khutyyaha ГУ,
Khevesyo
П.
In sedge, sedge-moss and moss-cotton
grass bogs, late snow areas, hummock dwarf
shrub sedge-lichen-moss and tussock dwarf
shrub Dicranum tundras, on sandy margin of
lake,
at the foot of Sphagnum tussocks on
margin of lake, usually on sandy soil, often
with Scapania hyperborea. Sporadically with
gemmae. For information about leaf
modifications that result from gemma
formation see Potemkin (1993b).
After investigation of Yamal collections of
Scapania tundrae, including some S.
pulcherrima-like phenotypes (mainly from
Khutyyaha, sedge-Sphagnum tussock bog,
July 30, 1977, E. N. Andrejeva, N 49) and
numerous transitional forms to S. tundrae
phenotypes, I think that S. pulcherrima,
described from West Greenland (Schuster
1974),
represents a form of S. tundrae,
developing in peculiar ecological conditions.
While Schuster (1974) considers that
Scapania pulcherrima "exhibits the same
relationships to S. degenii as does S. tundrae
to the S. irrigua-hyperborea complex", I can
not agree with this statement. S. pulcherrima
differs from S. degenii in its not persistent
and smaller oil-bodies; pigmentation and
broadly oval shape of its 2 - celled gemmae;
presence of coarse, irregular broad-based
teeth; ecological behaviour; etc. On the other
hand, all distinctive features of S.
pulcherrima or tendencies for their
manifestation occur in diverse phenotypes of
S. tundrae. S. pulcherrima shares with S.
tundrae cell size, oil-body and gemma
characteristics, and ecological behaviour.
Such features as short, winged and
semicirculary curved keel, minutely papillose-
verruculose cuticle and vinaceous
pigmentation of ventral leaf bases occur more
or less often also in Siberian plants of S.
tundrae (cf. Mueller 1951 - 58, Fig. 347: a,
b).
Schuster (I.c.) characterizes the leaves of
S. tundrae as entire-margined or bearing
scattered small teeth associated with gemma
formation. However, the tendency to develop
the crooked margin with solitary obscure
broad-based teeth or marginal projections not
associated with gemma formation is clear
from his Fig. 407. This feature is rather
common for Siberian plants of the species
and gives them a rather peculiar appearance.
It is not difficult to imagine the
transformation of such marginal projections
into irregular broad-based teeth of S.
pulcherrima.
Scapania paludicola Loeske & K. Muell.
var. rotundiloba Schust. (S. paludicola var.
paludicola auct. non Loeske & K. Muell. -
Gribova & Potemkin 1988)
AT:
Belyy I, Tambey I, Kharasavey II; NT:
Matyuiyaha
II, Bovanenkovo J, Tomboitosyo П,
Neromayaha
III, Mantyto I, Marre Sale 3, Khakhayayaha
II,
Lyakkotosyo I; ST: Khutyyaha II, Khevesyo IV,
Laptayaha
I, Kharangyneto 2, Er'yaha II, Khadytayaha
In sedge, Sphagnum, sedge-Sphagnum and
moss-cotton grass bogs, in moist dwarf
shrub-sedge-moss, grass-sedge, cotton grass
and dwarf shrub-5phgnum-cotton grass
tundras, near water courses and lakes, etc.
Rarely with androecia and gemmae, once
with perianth. Modifications of gemmiparous
leaves are like those in S. hyperborea
(Potemkin 1993b).
Variety rotundiloba is characterized by
obtuse to rotund edentate leaf lobes and
reddish gemmae. Vinaceous red pigmentation
of ventral leaf bases is usually absent in
Yamal plants. Only a few collections of the
variety were investigated in a fresh state,
and oil-bodies were usually (5)6 - 8(10) per
cell and 3-7 mkm in diam. The level of
genetic separation of var. rotundiloba from
the type variety is not clear and its
taxonomic status needs investigation.
Scapania cf. kaurinii Ryan.
ST:
Khevesyo I.
At the foot of peat outcrop in flat-centered
polygon with Calycularia laxa, Scapania
scandica, and Marchantia polymorpha.

The Hepaticae of the Yamal Peninsula 91
This report is based on sparse sterile
material. The plants are mod. parvifoUa-
colorata of the species. They are assigned to
Scapania kaurinii on the following criteria:
1) leaf lobes subequal, characteristically
dorsally arched and adaxially concave in
better developed plants; 2) keel distinct,
stiff,
2(3) - stratose throughout; 3) cortex 1 - 2 -
stratose, distinctly differentiated, of very
thick-walled cells; 4) trigones moderate,
straight-sided to faintly bulging. These
characters separate the plants under consi-
deration from all small phases of the
extremely variable S. hyperborea and from
other species of section Jrrigua (K. Muell.)
Buch that sometimes resemble S. kaurinii
Scapania uliginosa (Lindenb.) Dum. s. str.
AT:
Belyy I; NT: Neromayaha I; ST: Khutyyaha I,
Khevesyo I.
In grass-sedge, sedge-moss and cotton
grass sedge bogs usually with flowing water,
among species of Drepanocladus, Calliergon,
Sphagnum, Meesia, Paludella squarrosa,
Aulacomnium turgidum and associated with
Chiloscyphus fragilis, Gymnocolea inflata, G.
fascinifera, Cladopodiella fluitans, Scapania
hyperborea, etc. Once with androecia.
Yamal plants are not quite typical. Leaf
cells often develop prominent trigones and a
punctate-papillose to striolate cuticle at the
leaf base. Purple pigmentation occurs usually
only near the ventral leaf base.
Scapania subalpina (Nees ex Lindenb.)
Dum.
AT:
Belyy I (Zhukova & Rebristaya 1987).
In pool of sedge-moss bog.
Scapania crassiretis Bryhn
AT:
Kharasavey I; NT: Lyakkotosyo I; ST: Khevesyo
III.
In moist grass-sedge-dwarf shrub-moss,
dwarf shrub-lichen-moss, grass-cotton grass
and grass spotty tundras, more often on
moist bare ground, associated with Tri-
tomaria quinquedentata, Anastrophyllum
minutum, Barbilophozia kunzeana, B. bin-
steadii, Ptilidium ciliare, Cephalozia bi-
cuspidata, Scapania paludicola var. ro-
tundiloba, S. scandica, Odontoschisma ma-
counii, Lophozia ventricosa s.l., etc. Often
with gemmae. On bare soil Scapania
crassiretis often develops mod. densifolia-
colorata with leaves often rather briefly
decurrent ventrally. These plants can be
confused with S. degenii. A principal
difference of these species is different keel-
stem angles in the same insolation con-
ditions: in forms of S. crassiretis from
exposed sites the keel-stem angle in the
distal part of the keel does not exceed 110°,
but in S. degenii it is up to 160-185°; in
shade they are 80 - 90° and 100 - 120°
respectively (cf. Schuster 1974). Moreover
the keel of S. crassiretis is usually almost
straight in distal part vs more or less arched
in S. degenii; marginal teeth of S. crassiretis
are mostly uniseriate with a strongly
elongated apical cell (excepting the mod.
subintegrifolia-egemmipara which has weakly
developed, solitary, 1 - celled, slightly
elondated teeth) vs with slightly elongated
apical cell, often with two cells at base; cells
with coarse nodulose trigones in S. crassiretis
vs moderate, more or less bulging trigones in
S. degenii
Scapania simmonsii Bryhn & Kaal.
NT:
Yuribeitoyaha II.
On bare ground in herb-dwarf shrub moss
tundra, on hummock in sedge-willow-moss
hummock tundra, in moist depression in
hummock-pool sedge-yernik-willow tundra,
associated with Ptilidium ciliare, Ble-
pharostoma, Odontoschisma macounii, Ana-
strophyllum minutum, BarbUophozia
quad-
riloba f. glareosa, and Tritomaria quinque-
dentata.
Together with typical plants some weakly
developed shoots were found. They represent
mod. laxifolia-parvifolia-integrifolia-subcolo-
rata and mod. parvifolia-angustifolia-integri-
folia-colorata. Plants of the last modification
are only 0.6 mm wide and 2 mm long and
resemble very much the species of the sectio
Curtae (K. Muell) Buch. They have plane
leaves; cells with moderate, only sporadically
nodulose trigones; a comparatively weakly
papillose cuticle; and stem with a distinctly
differentiated, 1(2) - stratose, blackish cor-
tex of thick-walled cells.

92 A. D. POTEMKIN
DIPLOPHYLLUM (Dum.) Dum.
Diplophyllum taxifolium (Wahlenb.) Dum.
var. macrosticta Buch
AT: Belyy V, Kharasavey II; NT: Mantyto II; ST:
Khutyyaha I, Laptayaha Ш.
Predominantly in late snow areas mostly on
sandy, exceptionally on loamy, ground,
sporadic in some moss, moss-lichen, herb,
sedge and cotton grass tundras. According to
Zhukova & Rebristaya (1987) in Belyy the
species occurs in communities on sandy,
loamy and peaty ground. Gemmae rare.
The distribution of the species in Yamal is
not uniform: in many localities it is absent or
rare,
but in Belyy it is common species with
rather wide ecological amplitude.
GEOCALYCACEAE Klinggr.
Subfam. LOPHOCOLEOWEAE Rodway
CHILOSCYPHUS Corda
Subgen.
LOPHOCOLEA
(Dum.) ?Engel &
Schust.
Chiloscyphus minor (Nees) Engel &
Schust. (Lophocolea minor Nees)
NT: Neromayaha I; ST: Khytyyaha I.
In Sphagnum-sedge, herb-moss and lichen
willow stands on humus, associated with
Scapania curia, Cephalozia bicuspidata, C.
pleniceps, Blepharostoma, etc. With gemmae.
Plants from Neromayaha (lichen willow
stand) are very small to medium in size,
from 130 mkm to 1000-1500 mkm wide. The
smallest phases have an almost transverse
leaf orientation on the stem. Gemmae occur
in small quantity only on occasional plants.
They become free 2 - celled, more rarely in
3 - or 4 - celled more or less globose bodies.
Subgen. CHILOSCYPHUS
Although Chiloscyphus pallescens (Ehrh. ex
Hoffm.) Dum. was reported previously for
the peninsula (Andrejeva 1981; Zhukova &
Rebristaya 1986,1987; Gribova & Potemkin
1988)1 I have found no typical plants of this
species in investigated specimens. I consider
ail earlier reports doubtful also because of
the chromosome investigation of arctic
Chiloscypus (Steere & Inoue 1978), that
showed that large-celled Ch. pallescens-like
arctic phenotypes have 9, not 18
chromosomes and should be attributed to Ch.
polyanthos rather than to Ch. pallescens.
For differentiation of Chiloscyphus pal-
lescens s.l. and Ch. polyanthos s.l. oil-body
characteristics may be useful. According
to Schuster (1980) oil-bodies of Ch.
polyanthos are (1)2, more rarely 3-4(5) per
cell, large: 4 - 5 x 7 - 12 mkm to 6 - 7 x 8 -
12(18) mkm vs in Ch. pallescens mostly
3)4 - 6(10 - 12) to 8 - 12 per cell, smaller
spherical 4 - 5.5 mkm, oval 4 - 4.5(5) x 8 -
9(10-
11) mkm.
Unfortunately all Yamal specimens of
Chiloscyphus were studied without oil-bodies.
I retain Ch. fragilis as a species because most
Yamal specimens very good fit to it and
consequently almost no doubts were arisen
during the investigastion of the Yamal
materials. Nevertheless the status of this
species should be revised in the future on the
basis of not only morphological but
biochemical and chromosome characters.
Chiloscyphus fragilis (A.Roth) Schiffn. (Ch.
pallescens auct. non (Ehrh. ex Hoffm.) Dum.
- Andrejeva 1981; Zhukova & Rebristaya
1986,1987; Gribova & Potemkin 1988)
AT: Belyy I, Khabelyaha П, Tambey П, Tirvyyaha Л
Kharasavey II; NT: Matyuiyaha I, Bovanenkovo /,
Tomboitosyo II, Neromayaha III, Marre Sale //,
Yuribeitoyaha II, Khakhayayaha II; ST: Khutyyaha I,
Kheveeyo II, Er'yaha I.
Usually in sedge and cotton grass bogs,
moist sedge and cotton grass tundras, moss
willow stands, etc. Once with mature
capsules.
PLAGIOCHILACEAE (Joerg.) K. Muell.
PLAGIOCHILA (Dum.) Dum.
Plagiochila asplenioides (L.) Dum. subsp.
poreiloides (Torrey ex Nees) Schust. (incl. f.
subarctica (Joerg.) Schust.; P. arctica auct.
non Bryhn & Kaal. - Andrejeva 1981)
NT: Bovanenkovo 2; ST: Khutyyaha II.
In Equisetum-moss, moss and flood plain
willow stands, hummock moss-dwarf shrub
tundra, sedge bog; associated with
Drepanocladus uncinatus, Mesoptychia,
Lophozia gUlmanii, Aneura pinguis, Pellia
neesiana, etc.
ARNELLIACEAE Nakai
ARNELLIA
Lindb.

The Hepaticae of the Yamal Peninsula 93
Arnellia fennica (Gott.) Lindb,
NT:
Matyuiyaha I, Yuribeitoyaha Ш; ST: Khevesyo I,
Laptayaha
III, Kharangyneto /.
In diverse spotty tundras, herb-Equisetum
willow stands, old fire place, late snow area;
on bare and thinly turfed soil, associated
with Barbilophozia quadriloba, Lophozia
heterocolpos, L. grandiretis, Tritomaria sci-
tula, T. heterophylla, T. quinquedentata,
Scapania gymnostomophila, Jungermannia
polaris, Odontoschisma macounii, Crypto-
colea, Blepharostoma, Nardia geoscyphus,
etc.
The species varies from mod. densifolia-
colorata to mod. laxifolia-viridis.
CALYPOGEIACEAE (K, Muell.) H. Arn.
CALYPOGEIA Raddi
Calypogeia sphagnicola (H.Arn & J.Perss.)
Warnst. & Loeske
ST:
Laptayaha I.
In tussock swampy Dicranum-Sphagnum
tundra on tussock among Sphagnum with
Barbilophozia binsteadii.
Calypogeia muelleriana (Schiffn.) K.Muell.
NT:
Lyakkotoeyo I; ST: Khevesyo I, Laptayaha I,
Er'yaha
II.
In cotton grass and Ledum tundras, high-
centered polygon, moss-cotton grass bog,
associated mostly with Barbilophozia
binsteadii, Anastrophyllum minutum, Lopho-
zia ventricosa var. longiflora, Cep'iialozia
bicuspidata. Once (Er'yaha) with gemmae.
CEPHALOZIACEAE Migula
Subfam. CEPHALOZIOIDEAE
PLEUROCLADULA Grolle
Pleurocladula albescens (Hook.) Grolle. s.l.
AT:
Belyy III, Khabelyaha II, Tambey I, Kharasavey I;
NT:
Matyuiyaha II, Tiutey 4, Mantyto III, Khakhayayaha
III;
ST: Khutyyaha III, Khevesyo IV, Laptayaha II,
Er'yaha
II, Khadytayaha /.
Usually in late snow areas, rarely on spots
in spotty tundras. Once (Tiutey) with
gemmae (see Potemkin 1993b).
CEPHALOZIA (Dum.) Dum.
Cephalozia bicuspidata (L.) Dum. subsp.
ambigua (C. Mass.) Schust.
AT:
Belyy II (Zhukova & Rebristaya
1987),
Khabelyaha
I, Tambey I, Kharasavey I; NT: Tiutey /,
Tomboltosyo
I; ST: Khevesyo II, Laptayaha I, Er'yaha I.
In late snow areas and on spots in spotty
tundras. Sporadically with perianths, twice
with gemmae.
The overlapping of the variability ranges of
Cephalozia bicuspidata subsp. ambigua and
subsp. bicuspidata is considerable. Most
difficult and sometimes impossible to sepa-
rate is mod. parvifolia of the both subspecies.
All doubtful phenotypes of C. bicuspidata s.l.
were attributed to subsp. bicuspidata.
Schuster's (1988:180) arguments for distin-
guishing this taxon as a subspecies rather
than a species seem well founded.
Cephalozia bicuspidata subsp. bicuspidata
AT:
Belyy III, Khabeiyaha V, Tambey V, Kharasavey
III;
NT: Matyuiyaha III, Tiutey 2, Bovanenkovo 3,
Tomboltosyo
III, Neromayaha V, Mantyto III, Saletayaha
5,
Yuribeitoyaha I, Khakhayayaha V, Lyakkotoeyo II;
ST:
Khutyyaha V, Khevesyo V, Laptayaha Ш,
Kharangyneto
4, Er'yaha III, Khadytayaha 4.
In diverse moss, lichen, and spotty tundras,
late snow areas, herb formations on sand,
sedge-moss bogs, etc. Perianths and mature
capsules rather frequent, gemmae rare.
Cephalozia leucantha Spruce
NT:
Neromayaha I.
On humus-rich soil among crustose lichens
with Anastrophyllum minutum, Lophozia sp.,
in willow stand on loamy soil.
Cephalozia pleniceps (Aust.) lindb.
AT:
Belyy I, Khabeiyaha III, Tambey III, Kharasavey
I;
N1': Matyuiyaha I, Bovanenkovo 2, Tomboltosyo I,
Neromayaha
III, Mantyto П, Saletayaha 3, Yuribeitoyaha
II,
Khakhayayaha III, Lyakkotoeyo I; ST: Khutyyaha III,
Khevesyo
V, Laptayaha HI, Er'yaha HI.
On steep herb and nival slopes, in spotty,
mostly lichen-moss and sedge-moss, and
dwarf shrub sedge-(grass)-moss tundras, on
peat outcrops, edges of lakes and water
courses, in sedge-moss and moss bogs. Often
with perianths and mature capsules.
The species is very malleable. On exposed
bare soil it is represented by small, 1.5-4
mm long, very fleshy whitish phenotypes
with dense leaves and lobes terminated by
one obtuse not elongated cell. Perianths are
4 - 5 - stratose at the base and unistratose
only in upper
half,
occasionally only near the
mouth. In wet moss tufts comparatively large,
up to 27 mm long, phases of the species
occur. They are greenish, not fleshy, with

94 A. D. POTEMKIN
distant leaves with apiculate lobes of 1 - 2
cells.
Cephalozia Sunulifolia (Dum.) Dum.<? С
of finis auct. non Steph. Zhukova к
Rebristaya 1987 (the species was identified
in sterile state that is actually imposible (cf.
Schuster 1974)))
AT: Belyy I (Zhukova & Rebristaya 1987), Khrasavey
I; NT: Tomboltosyo I, Neromayaha I; ST: Khutyyaha III,
Khevesyo III, Khadyteyaha /.
In moist tussock moss tundras, Sphagnum
yerniks, peat outcrops, moss-lichen and
herb-moss tundras, sedge bog and willow
lichen nival tundra. Once (Khutyyaha) with
gemmae. Gemma formation leads to
development of reduced and multilobed
leaves as well as large underleaves (Potemkin
1993b).
The species is usually represented by forms
with distant, longly decurrent leaves. Most
vigorous plants may be to 40 mm long and 1
mm wide. In Kharasavey mod. densifolia was
found. The plants resemble Cephalozia
pleniceps. They have subtransverse oriented
shortly decurrent leaves with not connivent
lobes terminated usually by 1 cell. They
differ from C. pleniceps in the following
distinctive features: 1) antical leaf margin
decurrent in parallel to stem vs subtransverse
in C. pleniceps; 2) rhizoids often copiously
branched near the ends vs nonbranched,
dilated near the ends; 3) basal cell of
uniseriate lobe apicuius comparatively
narrow, ca. 28 - 35 mkm wide at the base vs
usually more than 33 mkm wide; 4) dorsal
cortical cells more than 3 times wider than
medullary cell diameter vs less than 3 times
wider in C. pleniceps; 5) ventral, stolon-like
branches heliotropic vs geotropic; 6) ali
plants sterile that is a characteristic feature
of the dioicous С lunulifolia, not of the
autoicous С pleniceps.
Some phases of Cephalozia lunulifolia
resemble the amphiatlantic species C.
macrostachya Kaal. They may develop
comparatively small cells in lobe bases;
leaves to 17-19 cells broad with
nonconnivent lobes; and similar stem
anatomy. They differ in the almost entire
perianth mouth and bract margins as well as
in thin-walled cells which are, at least partly,
more than 25 mkm wide in lobe bases.
Cephalozia connivens (Dicks.) Lindb.
NT: Neromayaha I.
In nival willow stand on bare slightly
humusy loamy sandy soil among Cephalozia
bicuspidata and C. pleniceps. Northernmost
report of the species.
Investigated plants differ distinctly from
admixed Cephalozia pleniceps in leaf
insertion; cell size (dorsal cortical cells 40 -
70 x 50 - 85 mkm vs 30(50) mkm wide; cells
of leaf lobe base 40 - 50 x 40 - 56 mkm vs
30,
rarely 35 - 40 mkm in diam.); leaves
only 9-14 cells wide with dorsal lobe 3-5
cells wide and ventral (3)4 - 5(6) cells wide
vs (14)16-23 cells wide with wider lobes
respectively; absence vs presence of stolons;
stem of 12-13 rows of cortical cells (5
dorsal rows and 7-8 lateral and ventral
rows) and 16-17 rows of medullary cells vs
24 rows of cortical and 32 rows of medullary
cells,
etc. The main difference from plants of
the temperate region is thin-walled medullary
cells.
Comparative study of temperate and
arctic species of Cephalozia as well as of
Jungermannia subgen. Plectocolea showed,
however, that development of thick-walled
medullary cells is characteristic only for
temperate populations and is considerably
rarer and more weakly expressed in the
Arctic.
Subfam. ODONTOSCHISMATOWEAE Buch ex
Grolle
ClADOPODIEUA Buch
Cladopodiella fluitans (Nees) Buch
ST: Khutyyaha II.
In bogs with flowing water among
Drepanocladus exannulatus, D. fluitans f.
pseudostramineum, Calliergon sarmentosum,
with Scapania uliginosa, S. hyperborea,
Gymnocolea fascinifera and among Sphag-
num with Gymnocolea fascinifera and G.
inflata var. acutiloba; in high-centered
polygon among Gymnocolea inflata var.
acutiloba.
The plants collected in bogs with flowing
water are quite typical but in the polygon
they are represented by mod. parvifolia-
fusca. They have a stem only 5 cells high,

The Hepaticae of the Yamal Peninsula 95
with distict underleaves and cortical cells
28 - 32 mkm wide and median leaf cells ca.
28 x 30 - 38 mkm.
ODONTOSCHISMA (Dum.) Dum.
Odontoschisma elongatum (Lindb.) Evans
NT: Matyuiyaha II, Mantyto H, Sebayaha I,
Khakhayayaha II; ST: Er'yaha I.
In diverse bogs, dwarf shrub sedge-lichen-
mpss and dwarf shrub-lichen tundras, on
steep slope, associated usually with Ptilidium
ciliare, Blepharostoma, Cephalozia bicus-
pidata, Gymnocolea inflata, Barbilophozia
binsteadii, B. kunzeana, Lophozia ventricosa
var. longiflora and Scapania irrigua, but on
steep slope also with Anthelia juratzkana,
Pleurocladula albescens, Calycularia laxa and
Anastrophyllum sphenoloboides.
The species varies from mod. densifolia-
colorata to mod. laxifolia-colorata vel
subcolorata, mod. viridis occurs very rarely.
Odontoschisma macounii (Aust.) Und.
AT: Belyy Ш, Khabeiyaha I. Kharasavey I; NT: Marre
Sale 1, Yuribeitoyaha II, Khakhayayaha I, Lyakkotosyo I;
ST: Khevesyo III, Laptayaha II, Er'yaha I, Kharangyneto
1.
On bare and thinly turfed soil in diverse
tundras, in flat centered polygons, moist
horse-tail nival and sedge willow stands,
apparently only in sites with comparatively
rich soils.
CEPHALOZIELLACEAE Douin
Cephaloziella divaricata (Sm.) Schiffn. var.
divaricata
AT: Khabeiyaha I, Tambey I, Kharasavey I; NT:
Matyuiyaha I, Tiutey 1, Bovanenkovo i, Tomboitosyo II,
Nea>mayaha II, Mantyto I, Marre Sale /, Yuribeitoyaha
II,
Khakhayayaha I, Lyakkotosyo I; ST: Khutyyaha II,
Khevesyo II, Laptayahs П, Kharangyneto 3, Er'yaha I.
In diverse moss, sedge, sedge-moss, dwarf
shrub and sedge-lichen tundras, cotton
grass-sedge-Spagnum bog, on peat outcrop,
on bare loamy ground on edge of temporary
water course. Often with gemmae, rare with
perianths.
Cephaloziella divaricata var. scabra (M. A.
Howe) S. Arn.
NT: Tomboitosyo I, Lyakkotosyo I; ST: Khevesyo I.
In sedge-moss and moss-herb willow
stands, hummock sedge-yernik moss tundra,
associated with Lophozia jurensis, L.
ventricosa, Ptilidium ciliare, Anastrophyllum
minutum, etc. Once with gemmae.
Cephaloziella divaricata var. polystra-
tosa (Schust. & Damsh.) Potemk. comb,
nov. (Cephaloziella byssacea (A. Roth)
Warnst. var. polystratosa Schust. & Damsh.,
Cladopodiella franciso (Hook.) Such ex
Joerg.
ST: Khutyyaha I, Khadytayaha 3.
In tussock tundras, on upland in bog and
on the margin of loamy landslip, associated
with Lophozia cf. savicziae, Gymnocolea
inflata, Cephalozia bicuspidata, Gymno-
mitrion corallioides, Scapania scandica, and
Marsupella sprucei. Often with gemmae,
perianths and mature capsules. Gemmae
more or less angulate, 1 - 2 - celled,
colourless to violetish, 23 - 25(39) mkm.
Spores from II - 13 to 15 - 16 mkm in diam.
Elaters 9-13 mkm in diam., with 2 brown
spirals 2.5 mkm wide. Two capsules found in
collection from Khutyyaha, margin of loamy
landslip, are not typical: valves coherent in
apical part of capsule; epidermal layer with
nodular thickenings on alterating longitudinal
walls;
and inner layer with mostly complete
semiannular bands.
CEPHALOZIELLA (Spruce) Schiffn.
Cephaloziella subdentata Warnst.
NT: Bovanenkovo 1, Tomboitosyo I, Neromayaha I,
Yuribeitoyaha I; ST: Khutyyaha II, Er'yaha I,
Khadytayaha /.
In willow-yernik sedge and sedge-Dryas-
lichen tundras, sedge-moss and polygonal
bogs,
associated with Sphagnum, Dicra-
num elongatum, Meesia, Blepharostoma,
Cephaloziella rubella subsp. arctogena,
Aneura cf. pinguis, etc. Often with perianths,
once with mature capsules and gemmae.
This species is often confused in the Arctic
with Cephaloziella rubella subsp. arctogena. It
differs from this taxon in sex distribution
(auticous vs paroicous), ability of hygric
forms to produce spinose teeth at leaf base as
in C. elachisla (Jack ex Gott. & Rabenh.)
Schiffn., and in perianth mouth, bract, and
capsule wall characters (see Potemkin
1992a).

96 A. D. POTEMKIN
0.5 mrn
Fig. 7.
Cephaloziella rubella
subsp.
arctogena.
(1) Young fertile shoot (x56). (2) Female bracts and bracteole (x80).
(3) Apex of female bract (хЗОО). (4) Sector of perianth mouth (хЗОО). (5) Median and basal cells of female bract
(хЗОО). (6,7) Male bracts (x80). (8) Leaf (x80). Scale bars: 0.5 mm - for 1; 0.4 mm - for 2, б - 8; 100 mkm - for 3 -
5. All from Khutyyaha (After Potemkin 1992a).
Phytologia, 63(5):327, 1987)
NT:
Neromayaha I, Saletayaha /.
On thinly turfed loamy sandy soil, associated
with Encalypta sp. With gemmae. First report
for Eurasian Arctic. Previously the variety
was known from South Greenland and, in
Eurasia, from Leningrad (St. Petersburg)
Province (for detailed descriptions and
illustratuions see Schuster 1988, Potemkin &
Tcherepanov 1993).
Cephaloziella rubella (Nees) Warnst. subsp.
rubella
NT:
Bovanenkovo 1, Marre Sale J.
In grass-sedge moss and moss-dwarf shrub
tundra. With perianths.
Cephaloziella rubella subsp. arctogena
(Schust.) Schust. & Damsh. (C. rubella auct.
non (Nees) Warnst. p.p. - Gribova &
Potemkin 1988; C. subdentata auct. non
Warnst. p.p. - Zhukova & Rebristaya 1986,
1987;
Gribova & Potemkin 1988) (Fig. 7)
AT:
Belyy I; NT: Matyuiyaha I, Bovanenkovo /,
Tomboltoeyo II, Neromayaha III, Mantyto I, Saletayaha
1,
Yuribeitoyaha II, Khakhayayaha II, Lyakkotosyo III;
ST:
Khutyyaha III, Khevesyo I, Kharangyneto /,
Khadytayaha 4.
In dwarf shrub-moss and sedge-moss
tundras, swampy sedge-(cotton grass)-moss
and dwarf shrub-moss (often Sphagnum)
communities. Often with perianths,
sporadically with mature capsules. For
differentiation see Potemkin 1992a).
Cephaloziella rubella cf. var. elegans
(Heeg.) Schust.
NT:
Tomboitosyo I, Marre Sale l\ ST: Khutyyaha I.

The Hepaticae of the Yamal Peninsula 97
In herb willow stands, spotty polygonal and
grass-sedge moss tundras, associated with
Cephaloziella rubella subsp. rubella and
subsp. arctogena, Scapania scandica, Lo~
phozia alboviridis, Cephalozia bicuspidaia, С.
pleniceps, and Anastrophyllum minutum. The
only one perianth have been seen. All Yamai
collections of this variety are poor. The
occurrence of Yamal plants together with the
other taxa of C. rubella leads to certain
doubts on their taxonomic isolation (cf.
Arnell 1956: 72).
Cephaloziella hampeana (Nees) Schiffn.
var. hampeana
AT:
Belyy I; NT: Matyuiyaha I, Bovanenkovo 1,
Tomboltoeyo II, Neromayaha II, Maiitytc I,
Khakhayayaha I; ST: Khevesyo П, Laptayaha II,
Khadytayaha 1.
In diverse moist sedge- and grass-moss
tundras, sedge and sedge-cotton grass bogs,
dwarf shrub-sedge-cotton grass and grass-
cotton grass tundras, sedge willow stands.
Sporadically with perianths and gemmae,
once (Khevesyo) with mature capsules.
Cephaloziella hampeana var. sibirica
C. Jens.
AT:
Belyy (Arnell 1918).
Cephaloziella arctica Bryhn & Douin (C.
rubella auct. non (Nees) Warnst. p.p. -
Zhukova & Rebristaya 1986; Potemkin 1989)
AT:
Belyy V, Khabeiyaha V, Tambey V, Kharasavey
Ш; NT: Matyuiyaha II, Tiutey 2, Bovanenkovo J,
Tomboltoeyo I, Neromayaha III, Yuribeitoyaha I,
Khakhayayaha II, Lyakkotosyo 11, ST: Khutyyaha II,
Khevesyo III, Laptayaha II, Er'yaha I, Khadytayaha 1.
In spotty, predominantly dwarf shrub-
sedge-(lichen)-moss, and swampy sedge-
moss tundras, at the foot of nival slopes, in
plant communities on weakly stabilized sand.
Sporadically with gemmae, perianths and
mature capsules.
In Arctic tundras, growing on bare soil, the
species usually develops mod. parvifolia-
colorata with purplish fuscous to purplish
black pigmentation; comparatively narrow
leaves, a little wider than the stem; and
reduced amphigastria, which are only
sporadically present near shoot apexes. In
moist moss tufts mod. viridis and mod.
subcolorata usually occur and often resemble
Cephaloziella hampeana. They differ from
the last species by the presence at least small
underleaves, thick-wailed cells of stem and
often of leaves, slightly elongated cells of
perianth mouth, female bract characters, etc.
Cephaloziella grimsulana (Jack ex Gott. &
Rabenh.) Lacout.
NT:
Neromayaha П, Sales&yaha 2.
On bare clay and humus clay soil on
landslip slopes, on loamy sand on steep
slope, associated with Barbilophozia hyper-
borean Lophozia excisa, etc. With gemmae,
once with juvenile perianth.
Cephaloziella uncinata Schust. (C. striatula
(CJens.) Douin var. argudeniata S. Arn. syn.
nov.; С subdentata auct. non Warnst. p.p. -
Andrejeva 1981; Zhukova & Rebristaya 1986,
1987)
AT I; NT: Matyuiyaha Ш, Tiutey J, Bovanenkovo 1,
Tomboitosyo I, Neromayaha ГУ, Mantyto I, Marre Sale 1,
Yuribeitoyaha II, Khakhayayaha II, Lyakkotosyo II; ST:
Khutyyaha II, Laptayaha I, Er'yaha II.
In diverse swampy communities of sedge,
cotton grass and mosses (often Sphagnum),
Sphagnum yerniks, dwarf shrub-(cotton
grass)-moss and herb-dwarf shrub-moss tun-
dras,
on margin of lake. Almost always with
perianths, sporadically with mature capsules.
The main peculiarity of Yamal plants is a
smooth or slightly papillose cuticle
characteristic not only for xeromorphic
phenotypes (Schuster 1980) but for mesic
and hygric forms also. This feature, however,
is very malleable in the species. Investigation
of Greenland material showed that the
character of the cuticle often varies
considerably from leaf to leaf on the same
shoot and from plant to plant in the same
specimen as in RMS 66 - 054, 66 - 306a,
66 - 1285a, 70 -2128, etc., and, con-
sequently, it is impossible to explain such va-
riability only by factor of moisture. The
other features of the species are quite typical.
Robust forms of the species may develop
solitary teeth, however, not only at leaf base
but on sinus sides also.
Investigation of the description, illustrations
and specimens of Cephaloziella siriatula var.
argudeniata from Novaya Zemiya (Arnell
1947),
which was assigned later by the
author (Arnell 1956) as a synonym of C,
subdentata var. spinigera H. Arn. & C. Jens.,

98 A. D. POTEMKIN
showed that it is almost identical with Yamal
plants of C, uncinata. It shares with this
species such important features as uncinate
leaf apexes of 1 - 5 superimposed cells with
strongly elongated terminal one, coarsely
dentate female bracts with uncinate apexes of
strongly elongated cells and a ciliate-dentate
perianth mouth of strongly elongated cells,
which are free on most of their length. The
only differences, excepting a faintly papillose
cuticle, are sporadically more narrow leaf
lobes,
only 4-7 cells broad at ba~ , and
longer uniseriate lobe apexes, to 5 cells long.
PTILIDIACEAE Klinggr.
PTILIDIUM Nees
Ptilidium clliare (L.) Hampe
AT V; NT V; ST V.
One of the most widespread Yamal hepatics
in diverse plant communities, excepting bogs
and dry lichen tundras on sand. Sporadically
it is a dominant species, forming large mats
about several hundred square metres.
Ptilidium ciliare is an extremely malleable
species. In Yamal it is mostly represented by
mod. densifolia of diverse coloration: from
green and yellowish green to brown, reddish
brown, deep purple and blackish violet. The
brown pigmented plants occur most often,
however. The number of marginal cilia varies
greatly also. There are often rather few cilia
in plants, growing on bare exposed soil
(mod. parvifolia ~ densifolia - colorata -oligo-
ciliata).
Sporadically forms with 25 - 40 cilia
in the ventral leaf margin occur. Their cilia
are composed of strongly elongated cells,
40-60x11-14(20) mkm, and they are
sometimes longer than the width of ventral
lobe.
Cells of these plants are usually larger,
than mentioned for the species, to 38 -
48(56) x (28)33 - 39 mkm at lobe base.
These differences are not very constant,
however, and such forms often grade into
typical ones,
ORDER MARCHANTIALES Limpr.
MARCHANTIACEAE (Bisch.) Lindley
MARCHANTIA L.
Marchantia alpestris (Nees)
Burgeff.
NT: Bovanenkovo 1, Saletayaha I.
On old fire place and on edge of water
course. With gemma receptacles, arche-
goniophores and antheridiophores.
Marchantia polymorpha L.
NT: Matyuiyaha I, Yuribeitoyaha II; ST: Khutyyaha
III, Khevesyo III, Er'yaha HI.
On edges of water courses, in karst funnels,
cotton grass bogs, at the foot of peat
outcrops, near and on fire places. Almost
always with gemma receptacles, arche-
goniophores and antheridiophores.
Marchantia aquatica (Nees)
Burgeff.
AT: Kharasavey I.
On margin of lake. With solitary gemma
receptacles.
PREISSIA
Corda
Preissia quadrata (Scop.) Nees subsp.
hyperborea Schust.
AT: Kharasavey I; NT: Neromayaha I, Lyakkotoeyo I;
ST: Khutyyaha III, Khevesyo I, Er'yaha I.
In crevices and on vertical surfaces of steep
sandy slopes, on river bank. Sporadically
with archegoniophores, antheridiophores and
mature capsules. Previously it was known
only from North America and Greenland
(Schuster 1992b).
The main distinctive features of this taxon
from the type subspecies, according Schuster
(I.c.),
are: plants normally autoecious (vs
dioecious); subarctic-arctic (vs predominantly
temperate-boreal); male receptacles with a
wide, thin, chartaceous, nitid flange, usually
arched upward (vs without such flange, the
margins not conspiciously curved upward);
carpocephalum with 2-4 sporophytes (vs to
8-10 sporophytes); spores smaller (ca. 50 -
70 mkm), with meshes of reticulations lower
(vs larger, ca. (62)65 - 80 mkm usually, with
meshes of reticulations high), etc.
Yamal plants of the subspecies are quite
typical but have smaller thalli, only 3-5 mm
wide (vs 10 - 15 mkm in original description)
that may be explained by unfavorable
conditions of soil nourishment (the other
basiphilous species in Yamal are smaller
also).
The spores are a little smaller too, 45 -
60 mkm.
RICCIACEAE Reichenb.
RiCClA L.
Riccia sorocarpa Bisch. subsp. arctica
Schust. {R.. sorocarpa auct. non Bisch. -

The Hepaticae of the Yamal Peninsula 99
Andre jeva 1981)
ST. Er'yaha I.
In willow-moss spotty tundra on loam with
Jungermannia polaris, J. obovata, Fossom-
bronia alaskana, Cephaloziella arctica,
Blepharostoma trichophyllum var. brevirete.
With mature capsules. Previously it was
known only from Greenland (Schuster
1992b).
AFONINA, О
ML
& i. DUDA [АФОНИНА, О М. & Й.
ДУДА) 1983. Contributions to the flora of hepatics
of Anadyr River Basin. - [Материалы к флоре
печеночных мхов бассейна реки Анадырь]. -
Novosti
Sist.
Nizsh. Rast. [Новости сист. низш.
рост./
20: 182-190.
ANDREJEVA, Е. N. [АНДРЕЕВА, Е. Н.] 1981. On
some main types of habitats of bryophytes in
southern hypoarctic tundras of Yamal. - (O
некоторых основных типах местообитаний
мохообразных в южных гипоарктических тундрах
Ямала].
In: Schljakov, R. N. (ed.)
BrioUkhenologicheskie issledovaniya vysokogornykh
raionov i Severa SSSR, Apatity [В кн.:
Бриолихенологические ' исследования
высокогорных районов и Севера СССР, (ред.
Шляков Р. Н.), Апатиты]. 18-19.
ARNELL, Н. W. 1918. Die Moose der Vega-Expedition.
- Ark. Bot. 15(5): 1-111.
ARNELL, S. 1947. Contributions to the knowledge of the
hepatics of Novaya Zemlya. - Svensk Bot. Tidskr.
41(2):
209-217.
ARNELL, S. 1956. Illustrated Moss Flora of
Fennoscandia. I. Hepaticae.
Lund:
Gleerups, 314.
CZERNYADJEVA, I. V. & A. D. POTEMKIN
[ЧЕРНЯДЬЕВА, И. В., А. Д. ПОТЕМКИН]
1993.
On the bryophyte flora of the Centra! Yamal.
- [К флоре мохообразных Центрального Ямала].
- Novosti
Sist.
Nizsh. Rast. [Новости сист. низш.
рост.] 29: 165-172.
DAMSHOLT, К. 1982. The perianth of Tritomaria
quinquedentata var. turgida (Lindb.) Weim.
(Hepaticae) - Bryologist &S(\):96-98.
GRIBOVA, S. A & A. D. POTEMKIN [ГРИБОВА, С.
А., А Д. ПОТЕМКИН] 1988. On the hepatic flora
This taxon, according to Schuster
(I.c.),
differs from the type subspecies in arctic
distribution, somewhat smaller spore size,
usually 65 - 86(91) mkm (in Yamal plants
(60)68 - 75(82) mkm); wing margins of
spores usually vestigial and locally developed
only or lacking; areolae tending to slightly
smaller, mostly ca. 5-8(10) mkm in diam
(in Yamal plants (4)5 - 8(9) mkm), etc.
of the Interfluve of Tomboy-Yakha and Se-Yakha
Rivers (the Central Yamal). - [К флоре
печеночных мхов междуречья рек Томбой-яха и
Се-яха (Центральный Ямал)]. Bot. Zhurn. [Бот.
журн.] 73(5): 685-690.
INOUE, Н. 1957. Notes on the taxonomical status of
Lophozia
diversiioba.
- Bot. Mag. (Tokyo) 70: 357-
362.
INOUE, H. 1961(1960). A new genus Hattorietla of the
Lophoziaceae. - /. Hattori Bot. Lab. 23: 37-40.
KONSTANTINOVA, N. A., A. D. POTEMKIN & R. N.
SCHLJAKOV 1992. Check-list of the Hepaticae and
Anthocerotae of the former USSR. - Arctoa 1(1-2):
87-127.
KOPONEN, Т., P. ISOVIITA & T. LAMMES 1977. The
bryophytes of Finland: an annotated checklist. -
Flora Fennica 6: 1-77.
LADYZHENSKAJA, K. I. {C. I.} [ЛАДЫЖЕНСКАЯ,
К. И.] 1971. New localities of a little-known species
Pseudolepicolea fryei (Perss.) Grolle & Ando in
Soviet Arctic. - [О новых местонахождениях
малоизвестного вида
Pseudolepicolea
fryei (Perss.)
Grolle & Ando в Советской Арктике]. Novosti
Sist.
Nizsh. Rast. [Новости сист. низш. рост.] 8:
314-317.
MACVICAR, S. M. 1926. The student's handbook of
British hepatics. 2nd ed. Eastbourne, 493.
MOGENSEN, G. S. & G. R. BRASSARD 1978.
Fossombronia alaskana found in Greenland. -
Bryologist 81(1): 155.
MUELLER, K. 1956. Die Lebermoose Europas (Musci
hepatici). Lf. 1-9. - In: Rabenhorsfs Kryptogamen-
Flora von
Deutschland,
Oesterreich u.d. Schweiz.
Leipzig, 6(3), 1365.
EXCLUDED TAXA
Erroneous identification Corrected identification
Barbilophozia
attenuata (Mart.) Loeske
Cephalozia affinis Steph.
Chiloscyphus pallescens (Ehrh. ex Eoffm.) Dum.
Plagiochila
arctica Bryhn & Kaal.
Scapania
brevicaulis
Tayl.
Scapania lingulata Buch
B.
binsteadii
C. lunulifolia
Ch.
fragilis
P.
asplenioides subsp. porelloides
S.
hyperborea
(S. helvetica phenotypes)
apparently a small form of Scapania sect. Irrigua
LITERATURE
CITED

100 A. D. POTEMKIN
PERSSON, H. 1946. Some Alaskan and Yukon
bryophytee. - Bryologist 49(2): 41-58.
POTEMKIN, A. D. [ПОТЕМКИН, А. Д.] 1988. On the
liverworts of the Yamal arctic tundras. - [O
печеночных мхах арктических тундр Ямала].
Труды 2 мал. конф.
ботаников
Ленинграда,
Ч. I.
Ленинград, апр. 1988. Л., АН СССР, Бот. Ин-т.
Деп.
ВИНИТИ 14.VII.1988, № 5682-В88 [Trudy
2 molod.
conf.
bot. Leningrada, Pt. I. Leningrad,
Apr. 1988. Leningrad, Akad. Nauk SSSR, Bot. Inst.
Msc Reserved VINIT1
14.
VII. 1988, № 5682-
B88] 1. 235-255.
POTEMKIN, A. D. {POTYOMKIN} [ПОТЕМКИН, А.
Д.] 1989. Liverworts of the Yamal arctic tundras.-
[Печеночные мхи арктических тундр Ямала].
Bot.
Zhurn.
[Бот. журн.] 74(6): 806-815.
POTEMKIN, A. D. [ПОТЕМКИН, А. Д.] 1990а. On
the variation of some hepatics from southern and
arctic tundras of the Yamal Peninsula. - [Об
изменчивости некоторых печеночных мхов из
южных и арктичесхих тундр полуострова Ямал].
Novosti Sist. Nizsh. Rast. [Новости сист. низш.
рост.] 27: 147-153.
[POTEMKIN, A. D.] ПОТЕМКИН, А. Д. 1990b.
Анализ модификационной изменчивости
печеночных мхов полуострова Ямал. - [Analysis
of modificational variability of hepatics of Yamal
Peninsula].
Труды З мал. конф. ботаников
Ленинграда, Ч. I. Ленинград, апр. 1990. Л., АН
СССР, Бот. Ин-т. Деп. ВИНИТИ 14. XI. 1990,
No 5700-B90 [Trudy 3 molod. con/, bot.
Leningrada,
Pt. 1. Leningrad, Apr. 1990. Leningrad,
Akad. Nauk SSSR, Bot. Inst. Msc Reserved
VINITI 14. XI. 1990, No 5700-B90] I: 235-255.
POTEMKIN, A. D. [ПОТЕМКИН, А. Д.] 1990с.
Lophozia alboviridis and L. decolorans (Hepaticae)
on the Yamal Peninsula. - [Lophozia alboviridis и
L decolorans (Hepaticae) на Ямале]. Bot. Zhurn.
[Бот.
журн.] 75(8): 1086-1092.
POTEMKIN, A. D. [ПОТЕМКИН, А. Д] 1990d. The
genus Tritomaria (Lophoziaceae, Hepaticae) in
Yamal. - [Род Tritomaria (Lophoziaceae,
Hepaticae) на Ямале]. Bot. Zhurn. [Бот. журн.]
75(12):
1742-1753.
POTEMKIN, A. D. 1990. The Uverworls of the Yamal
tundras. - In: Schljakov, R.N. (ed.) Abstr. 7th
Meeting
CEBWG.
Kirovsk, 54-55.
POTEMKIN, A. D. [ПОТЕМКИН, А. Д] 1991. On the
present state of understanding and prospects of
studying of the family Aneuraceae in the USSR. -
[О состоянии изученности и задачах по
изучению семейства Aneuraceae в СССР]. In:
Demkiv, ОТ. (ed.) Briologia v SSSR, ее
dostizheniya i perspektivy. Lvov, Akad. Nauk SSSR
& Akad. Nauk Ukr. SSR. [В кш Бриология в
СССР, ее достижения и перспективы (ред.
Демкив О.Т.), Львов, АН СССР, АН УССР]:
169-174.
POTEMKIN, A. D. [ПОТЕМКИН, А. Д.] 1992а.
Barbilophozia hyperborea (Schust.) Potemk. comb,
nov. and Cephaloziella rubella (Nees) Warnst. var.
arctogena Schust. in Sovet Arctic. - [Barbilophozia
hyperborea (Schust.) Potemk. comb. nov. и
Cephaloziella rubella (Nees) Warnst. var. arctogena
Schust. в Советской Арктике]. Novosti Sist. Nizsh.
Rast. [Новости
сист. низш.
раст] 28: 148-155.
POTEMKiN, A. D. 1992b. A new species of Prasardhus
(Hepaticae, Cymnomitriaceae) from the Yamal
Peninsula, West Siberian Arctic. - Ann. Bot. Fennici
29(4):
319-323.
POTEMKIN, A. D. [ПОТЕМКИН, А. Д.] 1993а. On
the infraspcclfic taxonomy, reproduction and
synonymy of Scapania scandica (H.Arn. et Buch)
Macv.(on the base of the Yamal materials). - [O
внутривидовой таксономии, размножении и
синонимике Scapania scandica (H. Am. et Buch)
Macv. (по ямальским материалам)]. Novosti Sist.
Nizsh.
Rast. [Новости сист. низш. раст.] 29:
153-157.
POTEMKIN, A. D. [ПОТЕМКИН, А. Д.] 1993b. On
the reproduction of some hepatics. - [O
размножении некоторых печеночников]. Novosti
Sist. Nizsh. Rast. [Новости сист. низш. раст.]
29:
145-152.
POTEMKIN, A. D. & I. V. TCHEREPANOV
[ПОТЕМКИН, А. Д., И. В. ЧЕРЕПАНОВ] 1993.
On the hepatic flora of the Leningrad Province. New
and littie-known taxa for the province. - [К флоре
печеночных мхов Лениградской области. Новые и
малоизвестные для облети таксоны]. Novosti Sist.
Nizsh.
Rast. [Новости сист. низш. рост.] 29:
158-165.
REBRISTAYA, О. V. [РЕБРИСТАЯ, О. В.] 1989.
Peculiarities of plant cover of the Yamal Peninsula. -
[Особенности растительного покрова полуострова
Ямал].
In: Shilov, V.N. (ed.) Kharakteristika
geologicheskikh i pochvenno-rastitetnykh osoben-
nostej territorij gazokondensatnykh mestorozhdenij
severa Tyumenskoj oblasti. Moscow. [В кн.:
Характеристика геологических и почленно-
растительных особенностей территорий
гаэоконденсатных месторождений севера
Тюменской области (ред. Шилов,
В.Н.),
М.,]:40-47.
SCHUAKOV, R. N. [ШЛЯКОВ, Р. Н] 1973.
Systematic notes on the family Lophoziaceae Cavers.
[Систематические заметки по семейству
Lophoziaceae Cavers]. Novosti Sist. Nizsh. Rast
[Новости
сист. низш.
раст.] 10: 287-302.
SCHUAKOV, R. N. [ШЛЯКОВ, Р. Н.] 1980-1981.
The hepatics of the North of the USSR. -
[Печеночные мхи Севера СССР]. Leningrad,
Nauka [Л, Наука] 3, 190 (1980); 4, 221 (1981).
SCHUSTER, R. M. 1969-1992. The Hepaticae and
Anthocerotae of North America east of the
hundredth meridian. - New
York-London,
Columbia
Univ. Press 2, 1062 (1969); 3, 880 (1974); 4, 1334
(1980);
- Chicago, Field Mus. Nat. Hist. 5, 854
(1992a); 6, 937 (1992b).
SCHUSTER, R. M. 1987. Studies on Metzgeriaies. I
North American Aneuraceae. - /. Hattori Bot. Lab.
62:
299-329.
SCHUSTER, R. M. 1988. The Hepaticae of South
Greenland. - Nova Hedwigia
Beih.
92: 1-255.
SCHUSTER, R. M. & K. DAMSHOLT 1974. The
Hepaticae of West Greenland from ca. 66 N to 72
N.
- Medd. Greenland 199(1): 1-373.
SCHUSTER, R. M., W. С STEERE & J. W. THOMSON
1959.
The terrestrial Cryptogams of Northern

The Hepaticae of the Yamal Peninsula 101
Ellesmere Island. - Natl Mas. Canada Bull 164: 1-
132.
SISKO, R. К. [СИСК0, P. K.] 1977. Introduction. -
[Введение].
In: Sisko, R.K. (ed.) Yamalo-
Gydanskaya
oblast'.
Leningrad: Gidrometeoizdat [B
кн.:
Ямало-Гыданская область (ред. Сиско,
Р.К),
Л.,
Гидрометеоиэдат]:
4-8.
STEERE, W. С. & Н. INOUE 1974. Fossombronia
alaskana, a new hepatic from Arctic Alaska. -
Bryologist 77(1): 63-71.
STEERE, W. С & H. INOUE 1978. The Hepaticae of
Arctic Alaska. - /. Hattori Bot. Lab. 44: 251-315.
YURTSEV, B. A., A. I. TOLMACHEV & 0. V.
REBRISTAYA [ЮРЦЕВ, Б. А., А. И. ТОЛМАЧЕВ
& О. В. РЕБРИСТАЯ] 1978. The Holistic
delimitation and subdivision of the Arctic. -
[Флористическое ограничение и разделение
Арктики].
In: Yurtsev, В.A. (ed.) Arkticheskaya
fhristicheskaya oblast'. Leningrad, Nauka. [В KW
Арктическая флористическая область (ред.
Юриев
Б.А.),
Л.: Наука] 9-67.
ZHUKOVA, A. L. & О. V. REBRISTAYA [ЖУКОВА,
А. Л. & О. В. РЕБРИСТАЯ] 1986. On the
liverwort flora of the Matyuiyakha River Region
(Yamal Peninsula). - [К флоре печеночных мхов
района реки Матюйяхи (полуостров Ямал)]. Bot.
Zhurn.
[Бот журн.] 71(5): 642-649.
ZHUKOVA, A. L. & О. V. REBRISTAYA
{REBRISTAJA} [ЖУКОВА, А. Л. & О. В.
РЕБРИСТАЯ] 1987. On the hepatic flora of Belyy
Island (Kara Sea). - [К флоре печеночных мхов
острова Белого (Карское море)]. Novosti Sist.
Nizsh.
Rast. [Новости сист. низш. рост.] 24:
208-213.
ZINOVIEVA, L. А. [ЗИНОВЬЕВА, Л. А.] 1969.
Sphenolobus cavifolius
(Buch & Am.) K. Muell. and
Sphenolobus minutus (Crantz) Steph. on the base of
the Ural materials (morphologic and systematic
notes).
-
[Sphenolobus
cavifolius (Buch & Am.) K.
Muell. и Sphenolobus minutus (Crantz) Steph. no
уральским материалам (морфологосистема-
тические заметки)]. Uchenye zapiski Permskogo
Gos. Univ. [Ученые записки Перм. Гос. Ун—та]
179:
282-288.