L. rupestris
Lewinskya
rupestris can be
recognized by combination of large plant size; immersed to emergent,
smooth capsules; exostome teeth when dry spreading at right angles
to capsule mouth; reduced endostome; and calyptrae with dense,
especially long, ascending hairs that mostly exceed the top of
calyptrae. Lewinskya
rupestris is a
temperate montane species present in most European countries;
northern and southern Africa; Central Asia; the Western Hemisphere
from Alaska to Bolivia; and SE Australia, Tasmania and New Zealand. In Russia the species is widespread in the Caucasus and southern
Siberia (Altai Mts, Sayans, Salair Range, Kodar Range) but somewhat
rare northward in the Kola Peninsula, Karelia, southern Urals,
subarctic eastern Siberia (Anabar Plateau), Yakutia (Orulgan Range)
and Chukotka. Although L. rupestris mostly
grows on siliceous rocks, in places where it is especially common it
also occurs on branches and twigs.
Capsules
short- or long-exserted 3
3.Endostome
segments erect when dry, filiform, smooth, ± half exostome length,
fragile, often broken off or absent 4
Endostome
segments incurved when dry, linear, papillose, as long as exostome
teeth, sturdy, usually persistent 5
4.Capsules
gradually tapered to setae; neck distinct; capsule slightly furrowed
in upper part 6. L. pylaisii
Lewinskya
pylaisii can be
recognized in field by its short-lanceolate leaves; short- to
somewhat long-exserted capsules that are slightly ribbed in upper
part and gradually narrowed to the setae; well-defined capsule
necks; 8 pairs of exostome teeth that cleft with age; exostome teeth
when dry spreading to reflexed at right angles to the capsule mouth;
and endostomes usually absent. Lewinskya
laevigata and L
pylaisii can be
confused because both are mostly saxicolous plants and have somewhat
short leaves, smooth, exserted capsules, and no endostomes. But, L. laevigata differs
from L. pylaisii
in capsule form (abruptly narrowed vs. gradually narrowed to setae);
exostome teeth number (8 pairs vs. 16 with teeth free or slightly
connate at base), and also never reflexed vs. varying from star-like
to reflexed. Lewinskya
pylaisii is a
widespread, amphioceanic Arctic/boreal species present in
Fennoscandia and the Baltic Sea islands, Iceland, Greenland,
Svalbard, Canadian Atlantic Provinces, Yukon, Subarctic Alaska, and
along the western coast of North America from the Aleutians to
northern California. In Russia this species has a bicentric
distribution: the coastal areas and islands of the Kola Peninsula
(Barents and White Seas) and Kamchatka Peninsula and Commander
Islands (Bering Island) in the Russian Far East. Lewinskya
pylaisii is a moss of
humid environments that grows on siliceous rocks near seashores.
Capsules
more or less abruptly narrowed to setae, neck absent; capsule smooth
in upper part 5. L. laevigata
Lewinskya
laevigata differs
from other species of Lewinskya,
as well as the saxicolous Orthotrichum
anomalum and Ulota
hutchinsiae in having
the following combination of features: exserted, somewhat short,
smooth capsules; usually 8-pairs of exostome teeth that are erect or
spreading at right angles to capsule mouth when dry; and endostome
mostly absent. For the differences between L. pylaisii and L. laevigata see
discussion under L. pylaisii. Lewinskya
laevigata has a
suboceanic holarctic distribution with localities in Europe (common
in Scandinavia and Iceland), throughout western North America, and a
few localities in temperate Asia. In Russia the species is similar
to L. pylaisii in
favoring humid, marine environments and having a bicentric
distribution: southern shores of the Kola Peninsula (Kandalaksha
State Reserve) and a few areas in the southern, temperate Russian
Far East. These southern Far East collections, although
morphologically close to L. laevigata appear
phytogeographically odd, and further study may find they represent a
cryptic species. Most records of Lewinskya
laevigata in northern
Asia (Anabar Plateau, Yakutia and Chukotka) represent either L. iwatsukii or a
putative, undescribed species of close to L. iwatuskii.
5.Leaves
blunt or acute; upper leaf cells with low, simple papillae[Ulota
hutchinsiae]
Leaves
acuminate; upper leaf cells with high, forked papillae 6
6.Capsules
smooth in upper part 4. L. iwatsukii
Lewinskya
iwatsukii differs
from all other Lewinskya
species in having the following combination of features: large
plants; narrow, lanceolate leaves; high forked leaf cell papillae;
exserted capsules; and exostome teeth spreading at right angles to
the capsule mouth when dry. In addition, L. iwatsukii differs
from other large, saxicolous Lewinskya
species in having
persistent endostomes that are ± as long as the exostome teeth, and
from corticolous species of Lewinskya,
e.g. L. elegans,
in having leaf cells with forked papillae. Lewinskya
iwatsukii was
originally described from Japan, but at present is known from the
Himalayas, India, Siberia, Russian Far East, the Caucasus and the
Polar Ural area. It is widespread and especially abundant in Arctic
East Siberia, Yakutia and Chukotka where it grows on basic and
calcareous sedimentary rocks, larch trunks, rocky soil, pebbly creek
bars, in open tundra communities, and occasionally on Populus,
Salix,
and Larix. Lewinskya iwatsukii
displays considerable variation in capsule morphology, particularly
in the the upper parts of the capsules. Our preliminary data on the
species indicate it may harbor one or many cryptic species.
Capsules
8-ribbed in upper part 4.
L. cf. iwatsukii(killiasiimorphotype)
Arctic
plants similar to L. iwatsukii, but with
capsules 8-ribbed distally are referred here tentatively to L. cf. iwatsukii,
killiasiimorphotype. Orthotrichum killiasii
Mull. Hal.,
described from Switzerland was accepted for Russian Arctic by
Abramova et al.
(1961), but later usually included in synonymy of L. speciosa. Preliminary
results of our molecular phylogenetic studies indicate that this
species might merit resurrection.
7(1). Capsules immersed to shortly emergent 7
Capsules
emergent or exserted 11
8.Capsules
smooth when dry 9
Capsules
ribbed when dry 10
9.Leaves
acuminate, rarely acute; exostome teeth 16; endostomial segments 16,
margins sinuose in outline 10. L. striata
Lewinskya
striata is
distinguished from other Lewinskya
species by the combination of deeply immersed, pale, smooth
capsules; 16 exostome teeth; and 16 fairly broad, lanceolate
endostomial segments. Curiously, sometimes specimens have emergent
capsules. Lewinskya
transcaucasica
differs from L. striata in having
shortly emergent rather than immersed capsules; 8-pairs of orange
rather than 16, pale exostome teeth; and having longitudinal crests
rather than papillae on the lower, inner (ventral) exostome surface. Lewinskya striata has
a holarctic distribution but is more common in western Eurasia and
western North America. Except for northern Scandinavia the species
is widespread in western Europe and also occurs in North Africa,
Turkey, Pakistan, China, Japan, and western North America from
southern Alaska to California. In Russia it is the most common
species of the genus in Kaliningrad Province and some parts of the
Caucasus. It also oρρurs sporadically in the Altai and is known
from a single locality in Volgograd Province). Lewinskya
striata occurs in
shaded, humid as well as exposed, dry habitats growing on Fagus,
Carpinus, Betula,
Padus, Salix and
Sorbus.
Leaves
rounded, obtuse or acute; exostome teeth 8-paired; endostomial
segments 8, margins entire in outline [L. vicaria]
Lewinskya
vicaria (Laz.) F. Lara, Garilleti & Goffinet was described from the Fergana Valley
in central Tian-Shan and is still known only from the type
specimens. This species was discussed by Lewinsky-Haapasaari (1994). Lewinskya vicaria
is similar to L. transcaucasica in
having 8 pairs of exostome teeth and 8, entire margined endostomial
segments. But, it differs in having obtuse to rounded leaf tips;
upper leaf margins distinctly crenulate due to protruding papillae;
and immersed capsules.
10.Capsules
below mouth red; endostome segments 16 2. L. dasymitria
Lewinskya
dasymitria is similar
to L. transcaucasica
inhaving shortly
emergent capsules; 8 pairs of irregularly split exostome teeth; and
16, entire-margined endostome segments. Itdiffers from L. transcaucasica as
well as all other species of the genus in having capsules that are
dark red and distinctly ribbed below the mouth. Somewhat similar
broadly rimmed capsules also occur in L. rupestris, but that
species differs from L. dasymitria in lacking
an endostome. This rare moss was described from Xizang (Tibet) and
is also known from from the Chinese provinces of Yunnan, Sichuan and
Shanxi. In Russia the species occurs at 450600 m elev. in the
Altai Mts near Teletskoe Lake. It grows on Sorbus
in fir forests and
Rhododendron twigs
in birch forests.
Capsules
concolorous throughout; endostome segments 8 1.
L. affinis
[fastigiatum
morphotype]
Lewinskya
affinis is here
considered in a broad sense to include L. affinis var. fastigiata. It differs
from L. affinis
s. str. in having smaller plants; capsule ribs (46 cells wide);
immersed, urceolate capsules that are distinctly contracted below
mouth; and sparsely hairy to smooth calyptrae. In contrast L. affinis s. str. has
larger plants; weaker capsule ribs; emergent, cylindric, capsules
not contracted below mouth; and moderately hairy calyptrae. In
Russia the L. affinis
var. fastigiata
morphotype occurs in Eastern European Russia (Belgorod Province) and
the Western Caucasus region.
11(7). Capsules emergent; setae equal to or shorter than capsules 12
Capsules
exserted; setae longer than capsules 17
12.Mature
capsules with 8 distinct, dark ribs and deeply furrowed along their
entire length 13
Mature
capsules smooth or with 8 narrow, short ribs, and not or slightly
furrowed above 14
13.Capsules
slightly emergent; exostome teeth not or slightly perforated along
median tooth line or fenestrate above 1. L. affinis
Lewinskya
affinis is closely
related to L. sordida
but differs in having mostly acute leaves; slightly emergent
capsules; paler exostome teeth not or slightly perforate along
median lines; and pale, naked or nearly naked, campanulate
calyptrae. In contrast L. sordida has acute to
acuminate leaves; strongly emergent to exserted capsules;
typically bright exostome teeth perforated along median lines and
fenestrate above; and golden, sparsely hairly, mostly conic
calyptrae. Lara et al.
(2009b) also found the two species differed in endostome
ornamentation: L. sordida weakly
ornamented, segments translucent; L. affine more strongly
ornamented with striae and papillae. In European Russia L. affinis has often
been confused with L. speciosa because both
species have strongly ribbed capsules. But, L. affinis differs from
the widespread L. speciosa in having
longer capsule ribs (extending nearly the length of the capsules);
shorter setae; and long-rostrate opercula. Lewinskya
affinis is widespread
in Europe and Macaronesia, it also occurs in northern and eastern
Africa, western and Middle Asia, north India, China, Japan,
temperate as well as continental areas of North America. In Russia
the species is restricted to European Russia (western provinces
mostly within the steppe zone) and the Caucasus. There are many
records of the species from more northern areas in the forest zone
(Murmansk Province, northeastern European Russia, and the Urals). But, in fact these collections represent L. speciosa or L. elegans. Records of
the species from the Russian Far East need further study before
their identity can be confirmed. Lewinskya
affinis grows on the
bark of deciduous trees.
Capsules
strongly emergent to exserted; exostome teeth perforated along
median lines and fenestrate above 8. L. sordida
Lewinskya
sordida is closely
related to L. affinis
but differs in setae length (capsules strongly emergent to exserted
vs. capsules slightly emergent) and exostome teeth form (fenestrate
above vs. entire above). However, at times (especially in the
Caucasus, where both species occur) they can be difficult to
separate (see comments under L. affinis). Lewinskya
sordida is sometimes
confused with L. elegans because both
have emergent to exserted capsules; reflexed exostome teeth; and
eight well-developed endostome segments. In L. elegans, however, the
capsules are longer, not contracted below mouth when dry, and the
endostome segments are pale, not translucent, and papillose. In
contrast L. sordida
has distinctly ribbed capsules that are contracted below mouth when
dry and translucent, smooth or nearly so endostome segments. Lewinskya sordida
is very polymorphous species and in fact likely represents the group
of related taxa. In particular, in southern part of Russian Far East
it varies in position of capsules, exostome teeth fenestration,
number of segments and calyptrae characters. In addition, a
Caucasian morphotype differs from Far Eastern ones. Furthermore, due
to underestimating variability of L. pylaisii, which has
parallel forms with L. sordida, its marginal
morphotypes are often referred to the latter species. Recently this
problem was essentially addressed by Vitt (2017) for Pacific Region,
but in european Arctic it is still actual. Lewinskya
sordida is primarily
a cool temperate to subarctic species distributed in eastern Eurasia
and eastern North America but it also occurs at higher latitudes
(Alaska, western Greenland, south Siberia) as well as the Caucasus. It has been reported from Arctic & Subarctic European localities
(Svalbard, Kola Peninsula) but these reports are in need of
confirmation. In eastern North America it is especially common in
New England and extends as far north as Labrador and Newfoundland;
in East Asia it is known from Japan, Korea, and China. In Russia L. sordida is the most
abundant Lewinskya
species (as well as
Orthotrichum
s.l. species) in the Russian Far Eas, an area dominated mostly by
the genus Ulota. In addition, it occurs in the mountains of South Siberia (Sayans and
Altai), and in the vicinity of LakeBaikal. This common forest zone
species is typically found in open forests or at forest edges in
relatively humid areas, including populated places. It grows on the
trunks/twigs of many trees and shrubs. Ignatov &
Lewinsky-Haapasaari (1994) found that in the Altai L. sordida and L. elegans were nearly
equally common and this appears to be case in the Eastern Sajan Mts. However, L. sordida
is gradually replacing L. elegans to the east.
14.Capsules
smooth 15
Capsules
ribbed above 16
15.Capsules
cylindric, exostome teeth in 8-pairs, never cleft; endostome
segments 8, margins smooth 4. L. elegans
Lewinskya
elegans is usually
treated as a synonym of L. speciosa in recent
European and Russian handbooks and check-lists. However, Vitt &
Darigo (1997) considered them distinct species. In northern European
Russia, east Caucasus and Siberia collections in this complex have
the smooth capsules that correspond to L. elegans, while most
plants from central European Russia have the distinctly 8-ribbed
capsules that correspond to L. speciosa s. str. Indeed, recent molecular phylogenetiρ study of this complex support
this view of the two taxa. Lewinskya
elegans differs from
other Russian species of Lewinskya
by following
combination of features: upper leaf cells with simple papillae;
capsules cylindric, smooth, emergent to short exserted; exostome
teeth in 8 pairs, reflexed, pale-orange; and endostome segments 8,
moderately wide, curved inward when dry, pale, papillose. The world
distribution of the species is still insufficiently known because of
the confusion between L. elegans and L. speciosa; in North
America it occurs between the 40th and 50th parallels in Canada in
the boreal and hemiboreal zones; in eastern North America it is
found in the Atlantic States and the Great Lakes area; and between
the 50th and 60th parallels in western North America, but not
extending to the Pacific coastal areas. In Russia L. elegans is widespread
and common in northern European Russia, northward to the southern
taiga subzone; the eastern Caucasus (in western Caucasus L. speciosa s. str. occurs); western, eastern and southern Siberia (except the Arctic),
and from scattered localities in the southern Russian Far East. In
the eastern Russian Far East it is gradually replaced by L. sordida.
Capsules
ovate-oblong; exostome teeth in 8-pairs, often cleft; endostome
segments 8 or 16, margins mostly irregularly beaded to subentire
11. L. transcaucasica
Morphologically,
L. transcaucasica
can be considered an intermediate species between L. striata and L. elegans because of
the presence of emergent to shortly exserted, smooth capsules;
exostome teeth mostly paired but not attached to one another; and
816 endostome segments. It can be separated from other Russian
Lewinskya
species by the following combination of features: capsules emergent
to exserted, shortly oblong or ovate, smooth or nearly so; and
exostome teeth orange, mostly in 8 cleft pairs with remarkable
crests on the inner (ventral) surfaces, 8 to 16, wide endostome
segments with bead-like or subentire margins. For the differences
between L. transcaucasica and L. vicaria/L. dasymitria see
disρussions under those species. Lewinskya
transcaucasica
was described from Georgia (Eckstein et
al., 2017) and is now
known from Dagestan, the Altai, Kotuyskoe Plateau (SE Taimyr),
Republic of Buryatia, Transbaikalia, Amur Province and the upper
Bureya River, Khabarovsk Territory (Fedosov et
al., 2017b). The
siberian collections of L. transcaucasica were
previously identified as L. striata. Lewinskya
transcaucsica is a
corticolous moss found on aspen, alder, birch, spruce, bird cherry,
etc. at moderately high (4001100 m) elevations. Both L. striata and L. transcaucasica occur
in the Altai area, but the first species inhabits moist, shady
forests, while L. transcaucasica occurs
in more xeric, exposed environments.
16.Capsules
concolorous throughout; endostome segments 8; widespread in European
Russia 9. L. speciosa
In
Russia L. speciosa
was until recently treated in a broad sense to include L. elegans. Both are
widespread, epiphytic species with exserted capsules, superficial
stomata and narrow endostome segments. Vitt & Darigo (1997)
recognized both as distinct species (see discussion under the latter
species), and their treatment is followed here. Lewinskya
speciosa differs fromL. sordida and L. affinis in having
shorter and less strongly differentiated exothecial ribs; pale
exostome teeth; and papillose, non-translucent endostome segments. It differs from L. iwatsukii in leaf
cell papillae form (simple vs. forked) and setae length (capsules
emergent to short-exserted vs. long-exserted). Lewinskya
speciosa is a
holarctic species mostly associated with cool, temperate and
hemiboreal zones; towards the southern end of its range it is a
montane species. In European Russia L. speciosa is the most
widespread, common species of the genus in broadleaved forests but
becomes rarer northward, in south taiga subzone, being substituted
by L. elegans,
and southward, in steppe zone. All specimens named L. speciosa from Siberia
and the Far East actually represent L. elegans or L.
cf. iwatsukii.Lewinskya speciosa is
a corticolous species often abundantly growing on most trees/shrubs
(including introduced ones) in middle European Russia, as well
as recently fallen logs, boulders, and concrete. The species seems
to prefer moderately shady conditions and when encountered
is usually found on multiple rather than single trees.
Capsules
dark, orange-brownish below mouth; endostome segments 16; rare
central Asian species2. L. dasymitria
17(11).Capsules
very long and narrowly cylindric, endostome segments nearly as wide
as exostome teeth, concolorous with them, bright-red 12. L. vladikavkana
Lewinskya
vladikavkana can be
recognized by the following combination of features: setae 23(4)
mm long; exceptionally narrow-cylindric, nearly smooth or ribbed
distally capsules; and mostly bright-red endostome segments that are
almost as broad as the exostome teeth. In Russia L. vladikavkana is
common at elevations up to 1000 m in the central and eastern
Caucasus from Kabardino-Balkaria to Dagestan (Ignatov et
al., 2010), and in
the northern Altai Mts. Plants from the Altai differ from Caucasian
plants in having somewhat larger spores (2429 vs. 2125 mm). Long considered a Russian endemic, L. vladikavkana is now
known from Kazakhstan (Hradilek et
al., 2011) and
northeast Turkey (Lara et
al., 2010). The
Chinese and Himalayan L. hookeri (Mitt.)
F.Lara, Garilleti & Goffinet is similar to L. vladikavkana in
peristome structure and in having exserted, narrowly cylindric
capsules, but differs in having larger spores, (25)3553 vs. 2129 mm,
and flexuose leaves with recurved margins. L. vladikavkana grows in
the same habitats as L. striata, L. elegans andL. sordida: on
trunks/twigs of Betula,
Padus,
Salix,
Abies,
Caragana
and Sambucus
at moderate elevations.
Capsules
wider, cylindric, ribbed or smooth; endostome segments much narrower
and distinctly paler than exostome teeth, pale orange or translucent
or endostome rudimentary 18
18.Leaves
short-lanceolate, acute; capsules gradually narrowed to setae; young
exostome teeth when dry spreading at right angles to capsule mouth,
later cleft into 16 teeth; endostome segments filiform, mostly
broken off 6. L. pylaisii
Leaves
lanceolate to linear-lanceolate, acuminate; capsules more or less
abruptly narrowed to setae; exostome teeth reflexed when dry, not
cleft when old or, if spreading at right angles to capsule mouth,
endostome segments broad, persistent 19
19.Capsules
urceolate, contracted below mouth when dry, with 8 strongly
developed ribs and narrow furrows; exostome teeth mostly orange to
red, strongly perforated above 8. L. sordida
Capsules
cylindric, not contracted below mouth when dry, smooth or slightly
ribbed above, with narrow ribs and wide furrows; exostome teeth pale
yellow, not or slightly perforated above 20
20. Capsules well exserted on rather long, (2)2.55 mm, setae;
mostly saxicolous 5
Capsules
shorther exserted, setae 1.52(2.5); corticoous 21
21.Capsules
8-ribbed above 9. L. speciosa
Capsules
smooth 3. L. elegans