U. coarctata
This
species is quite distinct in having, nearly smooth, pyriform
capsules that are strongly puckered at the mouth. Remarkably, this
odd feature occurs convergently in Orthotrichum
and Macromitrium. Additional important features of U. coarctata include:
long-acuminate, curved leaves; short, non-fused exostome teeth; and
mostly absent endostomial segments. In RussiaU. coarctata occurs
in Kaliningrad Province and at moderate elevations (530 to 1250 m)
in the Western Caucasus. It is a corticolous moss that grows on
beech and alder.
— Mature
capsules oblong-cylindrical, fusiform, urceolate or obconic,
8-ribbed throughout; capsule mouths evenly contracted 2
2.Dry
capsules fusiform, narrowed toward the mouth 3
— Dry
capsules oblong, cylindrical, urceolate or obconic, often strongly
contracted below mouth 5
3.Leaves
flexuose to loosely erect when dry; endostome absent6. U. drummondii
This
species can be distinguished in the field by the following
combination of features: curved to loosely erect leaves; fusiform
capsules gradually narrowed to the mouth and strongly furrowed above
resulting in irregularly angular to star-like capsule mouths; and
single, whitish peristomes with often fragile and broken exostome
teeth directed upward to spreading. In Russia U. drummondii is mainly
a Far Eastern species found in the humid, oceanic areas of Sakhalin
and Kuril Islands, as well as western and southern Kamchatka. In
some areas, e.g. Iturup Island (Kuril Islands), it is the most
common species of the family. There is a single collection of U. drummondii from one
of the Baltic islands in European Russia. Ulota
drummondii mostly
grows on willow, poplar and alder, often with other Ulota
species and/or Lewinskya
sordida.
— Leaves
crisped-contorted when dry; endostome present 4
4.Endostome
segments mostly smooth; spores 27–32 mm;
Asian species [U. delicata]
Ulota
delicata was
described from China (Fujian Province), and is also known from Japan
(Honshu). Although it has not yet been found in Russia the species
could occur in the southern Russian Far East. Indeed, some
collections from the coastal areas of Primorsky Territory and
Kamchatka have similar contorted to crisped leaves and fusiform
capsules. However, these collections have either young sporophytes
or old, eperistomate capsules and differ from U. delicata in having
acuminate rather than obtuse to acute leaves and smaller spores.
— Endostome
segments distinctly longitudinally striolate; spores 20–24 mm;
European species 1. U. bruchii
The
presence of fusiform capsules that are narrowed to the mouth,
distinguish U. bruchii
from U.
intermedia
and U. crispula.Ulota
bruchii differs from
U. crispa s. str. in having irregularly reflexed exostome teeth that are often
split in mature capsules. Garilleti et
al. (2000) indicate
the best way to separate U. bruchii from U. crispa s. lat. is the
presence of longitudinally striolate endostome segments.Ulota
bruchii is a European
Russia species found in Kaliningrad and Leningrad (Baltic Sea
islands) Provinces.
5(2). Basal juxtacostal cells rectangular, thin- to moderately
thick-walled, longitudinal walls straight, non- or weakly sinuose,
non-porose; leaves mostly flexuose to loosely erect 6
— Basal
juxtacostal cells elongate, strongly thick-walled, longitudinal
walls more or less sinuose, often porose; leaves straight, flexuose
to loosely erect, or crisped 10
6.Perichaetial
leaves differentiated; perichaetial leaf apices obtuse to rounded 7
— Perichaetial
leaves not differentiated; perichaetial leaf apices acute to
acuminate 8
7.Capsules
obconic to cylindric, rarer oblong, not contracted below mouth,
brown; exostome teeth 16, 380–510 mm
long 10. U. orientalis
Ulota
orientalis was
recently described from the Khabarovsk Territory (Sovetskaya Gavan);
it grows on alder trunks on steep, wind-blown, sea-facing slopes
(Fedosov & Ignatova, 2018). It differs from other Holoarctic
Ulota species
(except U. japonica,
U. pacifica and U. barclayi Mitt.) in
having small spores and somewhat thin-walled, rectangular, basal
juxtacostal leaf cells with longitudinal walls that are not sinuose
or porose. It is similar to U. megalospora Venturi
and U. rehmannii
in having obconic capsules that are gradually narrowed from the
mouth to the setae and striolate on the upper, inner surface of the
exostome teeth. Ulota
orientalis differs
from U. japonica in
having perichaetial leaves obtuse to rounded (vs. acute); capsules
brown, oblong to obconic not constricted below the mouth (vs. yellowish to light brownish, mostly urceolate, markedly constricted
below the mouth); bands of exothecial cells weakly differentiated
(vs. strongly differentiated); exostome teeth much longer, 16 (vs. shorter, mostly 8); and endostome segments slender, non-keeled,
composed of one cell row (vs. robust, keeled, composed of two cell
rows). For the distinctions between U. orientalis and U. pacifica see
discussion under the latter species.
— Capsules
urceolate, cylindrical to ovate, mostly contracted below mouths,
yellow to brownish; exostome teeth in 8 pairs, up to 350 mm
long [U. barclayi]
Vitt
(2014) considered this species closely related to U. japonica. Indeed, the
two species were treated as conspecific (Anderson et
al., 1990); however
Garilleti et al. (2015) and Caparros (2015) accepted it as a good species. It is a
western North American species described from Sitka Island, Alaska. Ulota barclayi
is similar to U. orientalis in having
differentiated perichaetial leaves, but differs from U. japonica, U. orientalis andU. pacifica in having
lightly colored capsules that are contracted below the mouth; 8
pairs of exostome teeth; and red rimmed, angled opercula. The
operculum characters are very useful in distinguishing U. barclayi from U. japonica when the
perichaetial leaves are poorly differentiated. According to Caparros
(2015) the species may occur in Asia.
8.Capsules
cylindrical to ovate, slightly inflated, not contracted below mouth,
weakly longitudinally ribbed; segments filiform11. U. pacifica
This
recently described species from the South Kurils (Iturup and
Shikotan Islands) grows on trees as well as coastal cliffs (Fedosov
& Ignatova, 2018). Ulota
pacifica is most
closely related to U. japonica, but differs
in having capsules cylindrical to ovate, slightly inflated, not
contracted below the mouth, and weakly ribbed (vs. mostly urceolate,
non-inflated, contracted below the mouth, and strongly ribbed);
endostome segments filiform, widened only in basalmost part
(vs. linear, gradually widened below); and leaf cells smooth or with
a few scattered papillae (vs. regularly papillose). Ulota
orientalis differs
from U. pacifica
in having remarkably differentiated perichaetial leaves and
typically turbinate capsules. Ulota
pacifica and U.
curvifolia
are saxicolous mosses that have somewhat short, often curved setae.
But, U. pacifica
differs from U.
curvifolia
in the following features: lower leaf cell thickness (thin-walled
vs. thick-walled); upper leaf cell papillae form (absent or sparse,
single vs. regularly present, mostly forked); mature capsules shape
(never urceolate vs. more or less urceolate); exothecial cell band
differentiation (weakly vs. strongly); and exostome teeth
ornamentation (papillose vs. distally ridged). The East Asian U. perbreviseta Dixon &
Sakurai resembles U. pacifica in having
short, curved setae; capsules not constricted below the mouth; and
filiform endostome segments. It differs in having crisped leaves and
thick-walled lower leaf cells.
— Capsules
urceolate, not inflated, contracted below mouths, strongly
longitudinally ribbed; segments linear 9
9.Perichaetial
leaves differentiated; opercula red rimmed at base, angled when
viewed from above [U. barclayi]
— Perichaetial
leaves not differentiated; opercula not rimmed at base, rounded when
viewed from above 9. U. japonica
Wang
& Jia (2012) considered U. japonica distinctive
in having somewhat small plants and rectangular, more or less
thin-walled basal juxtacostal leaf cells with non-sinuose,
non-porose longitudinal walls. Ulota
barclayi has all of
these features, but differs from U. japonica in the
following: perichaetial leaves (obtuse to rounded vs. acute);
capsules (oblong to obconic, brown vs. mostly urceolate, light
brown); exostome teeth number (16, not fused vs. 8 fused pairs). Some populations of U. japonica have weakly
differentiated basal leaf cells and these can be confused with U. reptans which is also
similar to U. japonica
in having somewhat
small plants, slightly curved leaves, papillose exostome teeth and
sparsely hairy calyptra, but its basal leaf cells are clearly
thick-walled.
Other distinctive differences between U.
japonica
and U. reptans
include: leaf size (1.6–2.7 vs. 1.0–1.5 mm
long); capsule mouths when dry (contracted vs. mostly not
contracted; and spore size (16–24 vs. 27–34 mm). Ulota japonica
is an amphipacific, oceanic, temperate and boreal species; in Russia
it is restricted to the Russian Far East: South Kurils, Sakhalin,
Primorsky/Khabarovsk Territories and Kamchatka. The species is
fairly common in the coastal areas of Primorsky Territory and the
South Kuril Islands. It grows on the trunks and branches of willow,
poplar, alder, spruce and fir in the forest belt, usually at low
elevations, but occasionally as high as the timber-line.
10(5). Plants saxicolous 11
— Plants
corticolous 12
11.Leaves
stout, mostly straight when dry; leaf cell papillae low, simple;
exostome teeth on dorsal (outer) surfaces papillose throughout
7. U. hutchinsiae
This
species is distinguished from all other Russian Ulota
species in having
saxicolous plants and leaves that are mostly straight when dry. It
differs from all Russian saxicolous species of Orthotrichum
in having superficial stomata. In addition its occurrence on acidic
rather than calcareous rocks distinguishes it from the saxicolous O. anomalum and O. urnigerum. The genus
Lewinskya
is similar to U. hutchinsiae in having
superficial stomata, but most Lewinskya
species are
corticolous or have smooth to weakly furrowed capsules with reduced
endostomes (e.g., L. laevigata and L. pylaisii). In
contrast, U. hutchinsiae has
distinctly furrowed capsules with well-developed endostomes. In
Russia U. hutchinsiae
is found in the western part of country: southern Karelia Republic,
Berezovij Island in the Baltic Sea, Leningrad Province, and the
Caucasus (Krasnodar Territory and North Ossetiya). The species is
also known from a highly disjunct locality along the middle course
of the Biya River in the Altai Mts (Bardunov, 1974; Ignatov &
Ochyra, 1994).
— Leaves
slender, contorted when dry; leaf cell papillae high, partially
forked; exostome teeth on upper dorsal (outer) surfaces
longitudinally ridged 5. U. curvifolia
Ulota
curvifolia can be
distinguished from all other Holarctic Ulota
species by the following combination of features: plants saxicolous;
leaves contorted when dry; leaf cell papillae high, partially
branched; capsules subspherical to short-urceolate; calyptrae
densely hairy; and exostome teeth longitudinally striolate on upper
dorsal (outer) surfaces. In addition, U. curvifolia has more a
northern distribution and/or occurs in higher elevations than all
other Ulota
species. It is a widespread species in Russia and the only Ulota
species found in the
subarctic and continental mountain areas wherever acidic crystalline
bedrocks occur. On the other hand it is unknown in boreal mountain
regions composed of basic to neutral rocks (Putorana Plateau) and
volcanic areas (Kamchatka). It is especially abundant on shaded
overhanging granitic and gneiss rocks where it usually forms
extensive mats; occasionally it occurs on meta-gabbro.
12.Leaves
strongly contorted or crisped when dry; calyptrae densely hairy 13
— Leaves
flexuose to contorted, calyptrae naked or sparsely hairy 17
13.Endostome
segments striolate below1. U. bruchii
— Endostome
segments mostly smooth below 14
14.Leaves
contorted to slightly crisped when dry; leaf bases scarcely widened,
weakly concave, elliptic, gradually narrowed to acumina; basal
marginal leaf cells weakly differentiated in 1–3 rows; endostome
segments uniseriate or with biseriate patches, transverse cell walls
thin4. U. crispula
This
species differs from U. crispa and U. intermedia in having
contorted to weakly crispate leaves; a weakly differentiated,
narrow, group of cells along the basal leaf margins; a different
exothecial cell band structure; and partly biseriate endostome
segments (see Caparros et
al., 2016). Ulota
crispula can be
confused with Ulota
species not related to U. crispa because of the
presence of leaves that are weakly widened at base and contorted
rather than crisped. It differs from U. japonica in
thick-walled basal leaf cells and mostly larger spores. It differs
from U. barclayi, U. rehmanniiandU. reptans in having
capsules that are contracted below the mouth and weakly
differentiated basal marginal leaf cells. In Asian Russia some
specimens are hard to place in either U. intermedia or U. crispula because they
are gametophytically similar to U. intermedia (strongly
crispate leaves; broad leaf bases; well-differentiated, broad band
of basal marginal leaf cells) but sporophytically agree better with
U. crispula
(exothecial cell
bands yellowish throughout; endostome segments fragile, partly
biseriate); in
addition, capsules in these plants are notably contracted below the
mouth. These difficult to place, intermediate collections are
widespread on the Kamchatka Peninsula and occur sporadically in the
Altai Mountains, Khabarovsk Territory (Botchi State Reserve), and
Far Eastern Pacific Islands (Sakhalin and Southern Kurils). In
Russia U. crispula
s. str. apparently
occurs only in the Caucasus. The status of these intermediate Asian
plants, as well as their relationship to U. crispula and U. intermedia needs to
be clarified in the course of an overall revision of the U. crispa complex in the
Pacific region.
— Leaves
strongly crisped when dry; leaf bases wide, concave, round-ovate,
abruptly narrowed to acumina; basal marginal leaf cells strongly
differentiated in (2–)3–9 rows; endostome segments uniseriate,
transverse cell walls thick 15
15.Mature
capsules very narrow, urceolate, strongly constricted below the
mouth when dry and empty; ribs separated by narrow furrows,
collapsed at the constricted area of the urn; exothecial cell bands
orange, 4–6 cell rows wi 3. U. crispa
Caparros
et al. (2016) distinguish this species from the closely related U. crispula and U. intermedia by the
presence of urceolate capsules that are strongly contracted below
the mouth; wide exothecial cell bands with narrow furrows in between
that collapse when the capsules constrict; and 8-paired exostome
teeth that never split into 16 teeth. From other Ulota
species it differs in having wide, concave leaf bases that abruptly
taper above; and strongly crisped leaves. Caparros et
al. (2016) consider
U. crispa
a temperate, amphioceanic species. It is widespread in southern and
Middle Europe, extending northward to southern Norway and eastward
to Turkey, the Caucasus and Transcarpathia. In East Asia the species
occurs in China, Japan, and Taiwan. In North America it occurs along
the Pacific coast (British Columbia and Washington) with a few
inland localities, but seems to be absent in Atlantic North America. In Russia the species occurs in the western part of the Russian
Caucasus where it is especially abundant in the southernmost Black
Sea coastal area, eastward to Kabardino-Balkaria; there are a
few specimens known from Primorsky Territory in the Russian Far
East. All specimens from middle European Russia, the Altai and other
parts of the Russian Far East previously identified as U
crispa are now
referred to U. intermedia or U.
cf. crispula.
— Mature
capsules wide, cylindrical to urceolate, not or weakly constricted
below mouths when dry and empty; ribs separated by more or less
broad furrows in the upper half of urn; exothecial cell bands
yellowish, 2–5 cell rows wid 16
16.Capsule
mouths reddish rimmed; exostome teeth mainly reddish with hyaline
border; endostome segments 16, filiform; spores 15–20(–23) m[U. longifolia]
Ulota
longifolia Dixon &
Sakurai is a Japanese endemic usually considered a subspecies of U. crispa. It differs
from U. crispa
s. str. in having capsule shape and coloration similar to U. intermedia (cylindrical,
not or slightly contracted below mouth, with light ribs, composed of
elongate cells);longer peristome
teeth (325–360 vs. 320 mm
long), and 16, filiform vs. mostly 8 (occasionally 16), linear
endostome segments. The relationships of U. longifolia are
difficult to evaluate because it exhibits a combination of features
found in U. intermedia
and U. crispa.
— Capsule
mouths not rimmed; exostome teeth pale throughout; endostome
segments mostly 8, broadly linear; spores 18–35 mU. intermedia
Ulota
intermedia differs
from all other Russian Ulota
species in having wide, rounded to obovate, notably concave leaf
bases that are abruptly narrowed to linear-lanceolate acumina;
leaves strongly crispate when dry; and obconic or short-cylindrical
capsules that are not contracted below the mouth. Other distinctive
features of the species include: wide basal leaf borders of short
rectangular cells; exothecial cell bands narrow, yellowish, with
incrassate transverse walls and hyaline lumina; and endostome cells
with thickened transverse walls (cf. Caparros et
al., 2016). Ulota
intermedia is an
amphioceanic species, but its exact distribution remains unknown
because for a long time it was considered conspecific with U. crispa. Ulota
intermedia and U. curvifolia are the
most widespread Ulota
species in Russia. In
fact, all specimens of the U. crispa complex in
Middle European Russia are U. intermedia;eastwardit extendsto Ryazan, Vladimir
and Kostroma Provinces. It is also present around Teletskoe Lake
(Altai Mts.) in places with mild, wet climatic conditions. Most
specimens previously named U. crispa from coastal
areas of the Russian Far East are also U. intermedia. It is a
corticolous species that grows on a wide array of trees: e.g.,
Chosenia, Populus,
Salix, Betula, Picea, Abies, Acer, and
Pinus pumila.
17(12). Leaves (1.4–)1.8–2.5(–2.8) mm long; upper/median leaf cells
weakly papillose to nearly smooth; basal marginal leaf cells
differentiated in 3–7 rows; exostome teeth weakly papillose or
smooth, always striolate on upper, inner (ventral) surfaces 12. U. rehmannii
Ulota
rehmannii can be
distinguished from most other Russian Ulota
species by the combination of curved leaves; strongly incrassate
basal leaf cells; conic, nearly smooth calyptrae; mature, old
capsules obconic; exostome teeth mostly fused in 8 pairs and
striolate on upper, inner (ventral) surfaces; and large (21–39 mm)
spores. It is similar to the East Asian U. reptans and western
North American U. megalospora Venturi
in having weakly curved leaves; cylindrical to obconic capsules;
large spores; and conic, nearly naked calyptrae. Ulota
rehmannii differs
from U. megalospora
in leaf apex form (acute to short-acuminate vs. long-acuminate),
stomata position (basal vs. median) and spore size (21–39 vs. 35–60 mm).
In Russia the species is fairly common in humid, hemiboreal mountain
forests from southern Siberia (Altai Mts) to the southern Russian
Far East (Sakhalin and Kuril Islands).
— Leaves
1.0–1.5(–2.0) mm long; upper/median leaf cells strongly
papillose; basal marginal leaf cells differentiated in 1–3 rows;
exostome teeth papillose on upper, inner (ventral) surfaces 13. U. reptans
Ulota
reptans is similar to
U. rehmannii
and U. megalospora in
having weakly curved leaves; cylindrical to obconic capsules; large
spores; and conic, nearly naked calyptrae. Wang & Jia (2012)
distinguished U. rehmanniifrom U. reptans on the basis
of its leaf size (1.5–2.8 vs. 1.0–1.5(–2.0) mm long); and
exostome teeth ornamentation (striolate vs. papillose). Ulota
reptans is known from
four Russian Far Eastern (Khabarovsk Territory and Sakhalin Island)
collections. In
addition, there are several anomalous specimens from Khabarovsk
Territory that exhibit the critical features of U. reptans in
transitional combinations. These collections require additional
study before they can be confidently placed. Ulota
reptans is a
corticolous species found in hemiboreal forests on spruce, fir and
alder.