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Journal of Bryology
ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/yjbr20
New national and regional bryophyte records, 66
L. T. Ellis, M. Alataş, M. Aleffi, W. R. Álvaro Alba, D. A. Becerra Infante, K.
A. Cárdenas Espinosa, Md N. Aziz, V. A. Bakalin, G. Bergamo Decarli, M.
Boiko, N. Zagorodniuk, L. M. Boiko, E. A. Borovichev, G. Brusa, M. J. Cano,
J. A. Jiménez, S. S. Choi, I. Draper, F. Lara, M. V. Dunlin, J. Enroth, T. Ezer, V.
E. Fedosov, E. Fuertes, R. Garilleti, B. Albertos, S. R. Gradstein, A. Graulich,
V. Hugonnot, C. W. Hyun, M. Kırmacı, F. Filiz, U. Çatak, N. A. Konstantinova,
A. N. Savchenko, M. Kropik, J. Kučera, H. Kürschner, E. Yu. Kuzmina, N.
S. Liksakova, D. Maity, P. Martin, T. T. McIntosh, H. M. H. van Melick, B.
Moncada, Cs. Németh, S. V. O’Leary, G. F. Peñaloza-Bojacá, S. A. Maciel-
Silva, S. Poponessi, A. Cogoni, R. D. Porley, A. D. Potemkin, M. Puglisi, S.
Sciandrello, K. K. Rawat, V. Sahu, R. R. Paul, M. Ryan, P. Saha, D. S. Salas, J. G.
Segarra-Moragues, F. Sguazzin, N. R. Shafigullina, J. R. Shevock, S. Ștefănuţ,
A. Uygur, S. Karaman Erkul, S. Ursavaş, A. Özen, H. G. Zechmeister & R. H.
Zander
To cite this article: L. T. Ellis, M. Alataş, M. Aleffi, W. R. Álvaro Alba, D. A. Becerra Infante, K. A.
Cárdenas Espinosa, Md N. Aziz, V. A. Bakalin, G. Bergamo Decarli, M. Boiko, N. Zagorodniuk,
L. M. Boiko, E. A. Borovichev, G. Brusa, M. J. Cano, J. A. Jiménez, S. S. Choi, I. Draper, F.
Lara, M. V. Dunlin, J. Enroth, T. Ezer, V. E. Fedosov, E. Fuertes, R. Garilleti, B. Albertos,
S. R. Gradstein, A. Graulich, V. Hugonnot, C. W. Hyun, M. Kırmacı, F. Filiz, U. Çatak, N. A.
Konstantinova, A. N. Savchenko, M. Kropik, J. Kučera, H. Kürschner, E. Yu. Kuzmina, N. S.
Liksakova, D. Maity, P. Martin, T. T. McIntosh, H. M. H. van Melick, B. Moncada, Cs. Németh, S.
V. O’Leary, G. F. Peñaloza-Bojacá, S. A. Maciel-Silva, S. Poponessi, A. Cogoni, R. D. Porley, A.
D. Potemkin, M. Puglisi, S. Sciandrello, K. K. Rawat, V. Sahu, R. R. Paul, M. Ryan, P. Saha, D.
S. Salas, J. G. Segarra-Moragues, F. Sguazzin, N. R. Shafigullina, J. R. Shevock, S. Ștefănuţ,
A. Uygur, S. Karaman Erkul, S. Ursavaş, A. Özen, H. G. Zechmeister & R. H. Zander (2021)
New national and regional bryophyte records, 66, Journal of Bryology, 43:2, 193-212, DOI:
10.1080/03736687.2021.1942590
To link to this article: https://doi.org/10.1080/03736687.2021.1942590
Published online: 16 Jul 2021.
Submit your article to this journal
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BRYOLOGICAL NOTE
New national and regional bryophyte records, 66
L. T. Ellis
a
, M. Alataş
b
, M. Aleffi
c
, W. R. Álvaro Alba
d
, D. A. Becerra Infante
d
, K. A. Cárdenas Espinosa
d
,
Md N. Aziz
e
, V. A. Bakalin
f
, G. Bergamo Decarli
g
, M. Boiko
h
, N. Zagorodniuk
h
, L. M. Boiko
i
,E. A. Borovichev
j
,
G. Brusa
k
,M. J. Cano
l
,J. A. Jiménez
l
,S. S. Choi
m
,I. Draper
n,o
,F. Lara
n,o
,M. V. Dunlin
p
,J. Enroth
q
,T. Ezer
r
,
V. E. Fedosov
s,t
,E. Fuertes
u
,R. Garilleti
v
,B. Albertos
v
,S. R. Gradstein
w
,A. Graulich
x
,V. Hugonnot
y
,
C. W. Hyun
z
,M. Kırmacı
aa
,F. Filiz
aa
,U. Çatak
aa
,N. A. Konstantinova
ab
,A. N. Savchenko
ab
,M. Kropik
ac
,
J. Kučera
ad
,H. Kürschner
ae
,E. Yu. Kuzmina
af
,N. S. Liksakova
af
,D. Maity
ag
,P. Martin
ah
,T. T. McIntosh
ai
,
H. M. H. van Melick
aj
,B. Moncada
ak
,Cs. Németh
al
,S. V. O’Leary
am
,G. F. Peñaloza-Bojacá
an
,S. A. Maciel-Silva
an
,
S. Poponessi
ao
,A. Cogoni
ao
,R. D. Porley
ap
,A. D. Potemkin
aq
,M. Puglisi
ar
,S. Sciandrello
ar
,K. K. Rawat
as
,
V. Sahu
as
,R. R. Paul
as
,M. Ryan
at
,P. Saha
au
,D. S. Salas
av
,J. G. Segarra-Moragues
aw
,F. Sguazzin
ax
,
N. R. Shafigullina
ay
,J. R. Shevock
az
,S. Ștefănuţ
ba
,A. Uygur
bb
,S. Karaman Erkul
bb
,S. Ursavaş
bc
,A. Özen
bc
,
H. G. Zechmeister
bd
and R. H. Zander
be
a
The Natural History Museum, Cromwell Road, London, SW7 5BD, UK;
b
Department of Plant and Animal Production, Munzur University,
Vocational School of Tunceli, Tunceli, Turkey;
c
International School of Advanced Studies, University of Camerino, Camerino (MC), Italy;
d
Grupo de Investigación Biología para la Conservación, Universidad Pedagógica y Tecnológica de Colombia, Colombia;
e
Botanical Survey
of India, Central National Herbarium, Howrah, India;
f
Botanical Garden-Institute, Russian Academy of Science, Vladivostok, Russia;
g
Viale
Rovereto, Trento, Italy;
h
Department of Botany, Kherson State University, Kherson, Ukraine;
i
Kherson Branch of Admiral Makarov
University, Kherson, Ukraine;
j
Institute of Industrial Ecology Problems of the North of the Kola Science Center of RAS, Apatity, Murmansk
Province, Russia;
k
University of Insubria, Varese, Italy;
l
Departamento de Biología Vegetal (Botánica), Facultad de Biología, Universidad de
Murcia, E-30100, Murcia, Spain;
m
Team of Ecological Survey Research, National Institute of Ecology, Seocheon, Korea;
n
Departamento de
Biología (Botánica), Facultad de Ciencias, Universidad Autónoma de Madrid, Madrid, Spain;
o
Centro de Investigación en Biodiversidad y
Cambio Global, Madrid, Spain;
p
Institute of Biology Komi Science Centre UB RAS, Kommunisticheskaja St., 28, Syktyvkar, 167982, Komi
Republic, Russia;
q
Botanical Museum, P.O. Box 7, FI-00014 University of Helsinki, Helsinki, Finland;
r
Department of Landscape Architecture,
Faculty of Architecture, Niğde Ömer Halisdemir University, Niğde, Turkey;
s
Geobotany Department, Lomonosov Moscow State University,
Moscow, Russia;
t
Botanical Garden-Institute, Far East Branch of Russian Academy of Sciences, Vladivostok, Russia;
u
Departamento de
Biodiversidad, Ecología y Evolución, Facultad de Biología, Universidad Complutense de Madrid, 28040 Madrid, Spain;
v
Departamento de
Botánica y Geología, Facultad de Farmacia, Universidad de Valencia, Avda, Vicente Andrés Estellés s/n, E-46100 Burjassot, Spain;
w
Department of Systematics, Biodiversity and Evolution of Plants, Albrecht von Haller Institute, University of Göttingen, 37073 Göttingen,
Germany;
x
Donceel, Belgium;
y
Conservatoire Botanique National du Massif Central, Chavaniac, Laffayette, France;
z
Plant Resources
Division, National Institute of Biological Resources, Incheon, Korea;
aa
Adnan Menderes Üniversitesi, Fen Edebiyat Fakültesi, Biyoloji
Bölümü, TR-09010 Kepez-Aydin, Turkey;
ab
Polar-Alpine Botanical Garden, Kola Science Centre, Russian Acadamy of Science, Kirovsk-6,
Murmansk Province 184256, Russia;
ac
Institute of Botany, University of Natural Resources and Life Sciences Vienna, Vienna, Austria;
ad
Department of Botany, University of South Bohemia, České Budějovice, Czech Republic;
ae
Freie Universität Berlin, Institut für Biologie,
Systematische Botanik und Pflanzengeographie, Altensteinstr, 6, D-14195 Berlin, Germany;
af
Komarov Botanical Institute of the Russian
Academy of Sciences, St. Petersburg, Russia;
ag
Department of Botany, University of Calcutta, Kolkata, India;
ah
Tetbury, Gloucestershire, UK;
ai
Department of Botany, University of British Columbia, Vancouver, British Columbia, Canada;
aj
Valkenswaard, The Netherlands;
ak
Licenciatura en Biología, Universidad Distrital Francisco José de Caldas, Torre de Laboratorios, Bogotá, Colombia;
al
MTA Centre for
Ecological Research, Institute of Ecology and Botany, Vácrátót, Hungary;
am
51 Wood Street, Wallingford, Oxfordshire, OX10 0AX, UK;
an
Departamento de Botânica, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, Brazil;
ao
Department of
Environmental and Life Science, University of Cagliari, Sardinia, Italy;
ap
Cera dos Pomares, Aljezur, Portugal;
aq
Komarov Botanical Institute
of the Russian Academy of Sciences, St. Petersburg, Russia;
ar
Department of Biological, Geological and Environmental Biology, University
of Catania, Catania, Italy;
as
CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow, India;
at
The Ministry of Forests, Lands,
Natural Resource Operations and Rural Development, B. C. Government, Victoria, British Columbia, Canada;
au
Department of Botany, Bijoy
Krishna Girls’College, Howrah, West Bengal, India;
av
Center for Biodiversity Research and Extension in Mindanao (CEBREM), Central
Mindanao University, Musuan, Bukidnon Province, 8710 Mindanao Island, Philippines;
aw
Departamento de Botánica y Geología, Facultad
de Ciencias Biológicas, Universitat de València, Avda. Dr. Moliner 50, E-46100, Burjassot, Valencia, Spain;
ax
Via Selvotta, Muzzana del
Turgnano, Udine, Italy;
ay
Institute of Environmental Sciences, Kazan Federal University, Kazan, Tatarstan Republic, Russia;
az
California
Academy of Sciences (CAS), 55 Music Concourse Drive, Golden Gate Park, San Francisco, CA 94118, USA;
ba
Institute of Biology Bucharest of
Romanian Academy, 296 Splaiul Independentei, 060031 Bucharest, P.O. Box 56-53, Romania;
bb
Department of Biology, Faculty of Science
and Letters, Aksaray University, Aksaray, Turkey;
bc
Department of Forest Engineering, Faculty of Forestry, ÇankırıKaratekin University,
Çankırı, Turkey;
bd
Institute of Botany and Biodiversity Research, University of Vienna, Vienna, Austria;
be
Missouri Botanical Garden, 4344
Shaw Boulavard, St. Louis, Missouri, 63110, USA
ARTICLE HISTORY First Published Online 16 July 2021
1. Anthoceros hispidus Steph.
Contributors. G. F. Peñaloza-Bojacá and S. A. Maciel-Silva
Brazil. (1) Truticultura Pousada e Restaurante
Lago Azul, Gonçalves, Minas Gerais, 22°42′34.2′′S,
45°56′39.8′′W, 1733 m a.s.l., along the main road, on
soil in an area liable to flood next to a stream, 20
November 2017, leg. Peñaloza-Bojacá, G. F. 1079
(BHCB N° 204399); (2) road from Fazenda Floresta
© British Bryological Society 2021
CONTACT L.T. Ellis l.ellis@nhm.ac.uk
JOURNAL OF BRYOLOGY
2021, VOL. 43, NO. 2, 193–212
https://doi.org/10.1080/03736687.2021.1942590
Published online 16 Jul 2021
Negra to Melhoramentos, Camanducaia, Minas Gerais,
22°48′52.8′′S, 46°05′34.7′′ W, 1252 m a.s.l., on soil at the
roadside of the Jaguarí River, 21 November 2017, leg.
Peñaloza-Bojacá, G. F. 1088 (BHCB 204400) and 1089
(BHCB 204401).
Anthoceros hispidus is currently recorded for Marti-
nique, Jamaica, Cuba, and Mexico, found growing on
soil in the rainforest at 1000–1200 m a.s.l. (Ibarra-
Morales et al. 2015). In Brazil A. hispidus was recorded
for the states of São Paulo and Rio de Janeiro (Hässel
de Menéndez 1990), but for several years it was con-
sidered by Gradstein and Costa (2003) as possibly con-
specific with Anthoceros punctatus L. However, recent
collections of hornworts in the state of Minas Gerais
studied with scanning microscopy have confirmed
the presence of A. hispidus in Brazil. Additional to the
previous records of this species in Brazil (São Paulo
and Rio de Janeiro), here we expand its distribution
range. Anthoceros hispidus is commonly found on soil
of exposed areas (e.g. roadsides) next to tropical sub-
montane and montane forests. It can be easily distin-
guished from A. punctatus by its spores, which have
their proximal surface with no foramen. Another
species frequently confused with A. hispidus, due to
its similar spores, is Anthoceros scariosus Austin.
However, these species can be easily distinguished
by the lack of dorsal lamellae in the gametophyte of
A. hispidus and the greater number of antheridia per
chamber (>10), whereas A. scariosus has abundant
dorsal lamellae and fewer antheridia per chamber
(<6; Ibarra-Morales et al. 2015).
2. Aschisma cuynetii (Bizot & R.B.Pierrot) J.Guerra &
M.J.Cano
Contributor. R. D. Porley
Portugal. Algarve: Vau, (ca. 700 m west of), 20 m a.s.l.,
37°07′07.39′′N, 8°34′00.06′′ W, on open base-rich
lichen-stabilised sandy ground overlying the barrocal
of a coastal clifftop, with sporophytes, 10 January
2021, leg. R. D. Porley s.n. (Hb. Porley).
This is the first confirmed report of Aschisma cuynetii
for Portugal. This Mediterranean species occurs as scat-
tered individuals or in compact tufts, and its exceed-
ingly small size, 1–2 mm tall, and ephemeral nature
may partly explain its apparent rarity. The present
population was mixed with Didymodon luridus
Hornsch. ex Spreng.; other Pottiaceous mosses that
grew nearby include Microbryum curvicollum (Hedw.)
R.H.Zander, M. starckeanum (Hedw.) R.H.Zander and
M. davallianum (Sm.) R.H.Zander. The small
unbranched shoots with oblong-lanceolate cucullate
leaves, loosely curled when dry, only weakly incurved
above and scarcely exceeding the sporophyte, partially
obscure a sessile cleistocarpous minutely apiculate
capsule, reminiscent in the field of a small sessile
Phascum Hedw. or Weissia Hedw. In Aschisma Lindb.,
however, whilst the upper laminal cells are papillose,
the leaf margin consists of one or two rows of
smooth cells. The cucullate leaf apex, wider nerve
and yellowish capsule at maturity distinguish it from
Aschisma carniolicum (F.Weber & D.Mohr) Lindb. A
European endemic, this species was hitherto only
known from Spain and the Balearics where it is
classified as Vulnerable. Aschisma cuynetii was erro-
neously reported from Portugal by Guerra (2006) but
was later excluded from the Mediterranean checklist
(Ros et al. 2013).
3. Bazzania imbricata (Mitt.) S.Hatt.
Contributors. V. A. Bakalin, S. S. Choi and C. W. Hyun
Republic of Korea. Jeollabuk-do, Jirisan National Park,
upper reaches of the Hansin Stream at N-facing slope of
the main Jirisan ridge. Pinus-Abies-broadleaved forest on
steep slope, open moist cliff, 35.31947°N, 127.69469°E,
1571 m a.s.l., 12 October 2019, leg.V.A.Bakalin&
S. S. Choi Kor-79-6-19 (VBGI), det. V. A. Bakalin
Bazzania imbricata is still a poorly understood
species with its core area in the Sino-Himalaya, from
where it was described. The morphological similarity
of B. imbricata to B. tricrenata (Wahlenb.) Lindb. has
led to a misunderstanding of the former taxon
outside of the Sino-Himalaya. Moreover, it was even
known as far eastward as Japan, under the name Baz-
zania fissifolia var. subsimplex Steph. ex S.Hatt., until
that name was treated as synonymous with
B. tricrenata (Zhou et al. 2012). However, Furuki
(2015) described the features differentiating these
two taxa and showed that B. imbricata indeed occurred
in Japan. Within Japan this taxon represents the
floristic connections with the Sino-Himalaya. The pres-
ence of the species in the Korean Peninsula reflects the
same phytogeographic phenomenon and is, probably,
the link joining the Japanese and Sino-Himalayan
populations of the taxon. The fundamental feature
that helps to distinguish B. imbricata from
B. tricrenata is the surface of the oil body, which is
finely granulate in the former, and completely
smooth to loosely segmented in the latter. Another
feature is the comparative width of the underleaves.
Those of B. imbricata are commonly 3–4 times as
wide as the stem and have a very shallowly lobulate
apex, versus underleaves commonly 2–3 times wider
than the stem and more prominently lobed in
B. tricrenata. However, as noted by Furuki (2015) and
confirmed by our observations, the underleaf charac-
ters are subject to great variation and cannot always
help to distinguish the two species in their poorly
developed phases.
4. Delongia glacialis (C.C.Towns.) N.E.Bell, Kariyawa-
sam, Hedd. & Hyvönen
Contributors. K. K. Rawat, V. Sahu and R. R. Paul
India. Arunachal Pradesh, Tawang, Se La Pass near
lake, 27°30′47.1′′N, 92°05′45.7′′ E, 4120 m a.s.l., on soil,
194 L. T. ELLIS ET AL.
17 June 2015, leg. K. K. Rawat (LWG 300255F), same
locality, 14 November 2019, leg. K. K. Rawat and
R. R. Paul (LWG 325657D).
Delongia glacialis is reported for the first time from
Arunachal Pradesh, India. This species was previously
reported from Kashmir and Uttarakhand in the
western Himalaya (Townsend 1998,asOligotrichum
glaciale C.C.Towns.; Ellis, Alataş, et al. 2017). The
plants were blackish-brown, erect, and unbranched,
8–15 mm high, leaves closely imbricate, crowded at
apex, curled and appressed to stem when dry, deeply
concave, ovate-lanceolate, 1.8–2 mm long, 0.8–1mm
wide, apex acute, margin undifferentiated, dentate
and revolute at apex. The leaf cross-section has
adaxial lamellae 4–9 cells high, confined to midrib
only, abaxial lamellae absent, end cells of lamellae
not differentiated; costa 140–160 µm wide at base,
ending below the leaf apex; apical leaf cells thick-
walled, short quadrate, 12–20 µm wide, basal cells
thin-walled, rectangular, 20–60 µm long and 12–16
µm wide. Sporophytes not observed.
5. Didymodon eckeliae R.H.Zander
Contributors. M. J. Cano and J. A. Jiménez
Greece. Crete, Chania, near Samaria Gorge, 1200 m
a.s.l., 35°18′N, 23°55′E, 22 June 2018, leg. M. J. Cano
et al. 10341, 10342, 10347 (MUB, NHMC).
Didymodon eckeliae is a disjunct species with popu-
lations found in British Columbia, California, Nevada,
and Oregon along the western coast of North America
(Zander 2001,2007;Brindaetal.2014;Kofranek2014),
and the Iberian Peninsula in Europe (Puche et al. 2006;
Jiménez and Cano 2012; Ellis, Afonina, et al. 2014).
Recently, the species has been reported from Cyprus
(Cano 2020), with its range extending to Asia. During a
botanical expedition to Crete, a small population of
D.eckeliae was found growing on soil banks near a trail
path together with Syntrichia echinata (Schiffn.) Herrnst.
&Ben-Sasson,andTortula subulata Hedw. The new
record here reported from Crete confirms the distribution
of this taxon in the eastern Mediterranean area and is
also the first report from Greece. The specimens have
no sporophytes. Important diagnostic features of the
species include the regular bistratose leaf margins that
are broadly sinuose-crenulate, long-lanceolate leaves
and long-acuminate apices.
6. Didymodon tophaceus subsp. sicculus (M.J.Cano,
Ros, García-Zam. & J.Guerra) Jan Kučera
Contributors. J. Kučera and V. E. Fedosov
Russia. Eastern Caucasus, Dagestan Republic, Gunib
District, at the edge of the road from Gunib Town to
the field station of Gorny Botanical Garden of RAS,
42.38821°N, 46.95789°E, ca. 1330 m a.s.l. on the base
of concrete construction together with Tortula
muralis Hedw., 26 May 2010, leg. V. E. Fedosov,
MW:9068061 (admixture in specimen of T. muralis).
Didymodon tophaceus subsp. sicculus, whose taxo-
nomic status was recently investigated by Kučera
et al. (2018), has so far been reported from 18 Euro-
pean countries and Algeria (Jiménez et al. 2004; Hod-
getts and Lockhart 2020). Although sporadic records
were reported from northwestern and northeastern
Europe (The Netherlands and Estonia), the distribution
focus of this taxon is clearly in Mediterranean
countries. The new locality extends its known distri-
bution eastwards to an area which supports numerous
rare xerophytic species of predominantly Mediterra-
nean distribution These include, among others, Cross-
idium squamiferum (Viv.) Jur. and Tortula atrovirens
(Sm.) Lindb., which co-occur with Asian xerophytic
elements such as Entosthodon handelii (Schiffn.) Laz.
and Indusiella thianschanica Broth. & Müll.Hal.
(Ignatov et al. 2010).
7. Diplophyllum albicans (L.) Dumort.
Contributors. N. A. Konstantinova and A. N. Savchenko
Russia. Franz Josef Land Archipelago, Land Alexandra
Island, Severnaya Bay, remnant of the basalt plateau
(ca. 20 m high) with a large-column separation with
prismatic columns, 80°46′37′′N, 47°48′55′′ E, 22 m
a.s.l., in crevices and on ledges on soils and peat in
cushions among mosses and lichens, scattered on
edge of mats with dominant Marsupella boeckii
(Aust.) Lindb. and Gymnomitrion corallioides Nees and
admixture of Trilophozia quinquedentata (Huds.)
Bakalin, Gymnomitrion concinnatum (Lightf.) Corda,
12 July 2019, leg. A. N. Savchenko, CA16-19-6a
[KPABG(H): 123005, kpabg.ru›cris/?q=node/16].
The plants have a quite unusual appearance. They
are red brown, very small at 0.7 mm wide and 2–3
mm long, which is much smaller than the size indi-
cated in the literature (Schuster 1974; Damsholt
2002). Characteristic for the species, the leaf lobes
have elongated median cells and small thick-walled
marginal cells; one plant in the specimen has rotun-
date-stellate gemmae.
Diplophyllum albicans is not rare in the arctic-
montane Holarctic. This locality in Franz Josef Land is
about 60 km north of that in Prins Oscar Land (Nordau-
landat, Svalbard), which was previously considered the
northernmost locality of the species (Konstantinova
and Savchenko 2008).
8. Eobryum anoectangioides (Müll.Hal.) R.H.Zander &
P.Sollman
Contributor. R. H. Zander
Papua New Guinea. Morobe Prov., Mt. Sarawaket
southern range, 4 km NEE of Lake Gwam, open grass-
land (kunai) with scattered tree ferns, moist, diffusely
lit limestone cliff, 2850 m a.s.l., 147°9.5′E, 6°19′S, July
4, 1981, leg. D. H. Norris 62387 (MO).
New to the southwestern Pacific Islands, Eobryum
anoectangioides (Pottiaceae) was identified in the
JOURNAL OF BRYOLOGY 195
herbarium as Hymenostylium recurvirostrum (Hedw.)
Dixon and an unpublished forma name. This rare and
putatively ancient species is previously known from
two collections in India and one in Myanmar (Zander
and Sollman 2020). The genus Eobryum R.H.Zander &
P.Sollman is similar to Ardeuma R.H.Zander & Hedd.,
the latter a replacement name for all species of Hyme-
nostylium Brid. except the type, H. xanthocarpum
(Hook.) Brid. Eobryum differs in having a stem with
central strand, cauline leaves little tapering distally,
base not widened or somewhat narrower than
midleaf, and costal section with distinct epidermal
layers. The specimen was sterile.
9. Fossombronia delgadilloana Crand-Stotl., Stotler &
J.C.Benavides
Contributor. S. R. Gradstein
Ecuador. (1) Napo province, Páramo de Loma Gorda,
along Río Antisana below Micacocha, 78°10′W, 00°
37′S, in wetland with dominance of cushion plants
and Cortaderia Stapf, 3850 m a.s.l., 3 November 1979,
leg. L. B. Holm-Nielsen 20811 (GOET); (2) Pichincha pro-
vince, Páramo de Guamani, ca. 5 km W of Paso de la
Virgen, 78°10′W, 00°37′S, partly burned Polylepis Ruiz
& Pav. forest, in running water, deeply shaded, 3750 m
a.s.l., 8 February 1984, leg. S. Lægaard 51365 (GOET).
Fossombronia delgadilloana is a robust, semi-aquatic
species that was recently described from Mexico and
the high Andes of Venezuela where it grows submerged
in tropic-alpine mires at elevations between 3100 and
3900 m (Crandall-Stotler et al. 2019). The species is
recorded here as new to Ecuador; the two Ecuadorian
localities constitute the southernmost records of the
species. Fossombronia delgadilloana is closely related
to F. peruviana Gottsche & Hampe (= Austrofossombro-
nia peruviana [Gottsche & Hampe] Crand.-Stotl. Stotler
& A.V.Freire), a characteristic liverwort of mires in
páramo and humid puna vegetation in the high
Andes, from Venezuela to northern Chile, and resem-
bling F. delgadilloana in size, growth form and habitat.
Fossombronia delgadilloana, however, lacks any second-
ary pigmentation and its leaves are much thinner than
those of F. peruviana, mostly unistratose (except near
the base) and planate. Plants of F. peruviana, in contrast,
are usually reddish or brownish pigmented and have
rather fleshy, crispate leaves that are broadly pleated.
Moreover, the two species have different spores. A mol-
ecular analysis recovered the two species as separate
lineages (Crandall-Stotler et al. 2019). Further occur-
rences of F. delgadilloana in the Andes may be
expected.
10. Harpalejeunea grandis Grolle & M.E.Reiner
Contributor. S. R. Gradstein
Ecuador. Napo province, NE-facing ridge on the N side
of Cerro Sumaco, virgin moist páramo with scattered
shrubs, on steep slope, 77°39′W, 00°35′S, epiphyte on
Blechnum L. trunks, 3600–3700 m a.s.l., 24 April 1979,
leg. B. Løjtnant & U. Molau 12696 (GOET).
Harpalejeunea grandis is a rare northern Andean
species known from a few localities in the Eastern Cor-
dillera of Colombia (Grolle and Reiner-Drehwald 1999).
The species grows on bark and humic soil in páramo,
at about 3200–3800 m. The record from Ecuador consti-
tutes the southernmost locality of the species and the
first one from Ecuador. The species is readily recognised
by its relatively large size, with leafy stems 1–2mmwide
and to 3 cm long, resembling a laxly growing Lejeunea
Lib., and 4 cells wide ventral merophytes on main
shoots. Moreover, the underleaves are relatively large,
3–4× stem width, with an undivided to shallowly
retuse apex, and the ocelli in leaf lobes are arranged
in 1–2 unbroken rows. Because of its unusual morpho-
logical features, the species has been placed in a separ-
ate subgenus, H. subg.Cleefiolejeunea Grolle &
M.E.Reiner (Grolle and Reiner-Drehwald 1999).
11. Lejeunea suffruticola Spruce
Contributor. S. R. Gradstein
Ecuador. Galápagos Islands, Cerro Azul, on horizontal
branches of Pisonia floribunda Hook.f. in semi-decid-
uous Pisonia-Hippomane woodland, 250 m, 21 June
1976, leg. S. R. Gradstein H375 (GOET, U).
Lejeunea suffruticola is a very rare species that is only
known from the type specimen collected by Richard
Spruce over 150 years ago in the Andes of Ecuador
(Guataxi, 2200 m, on small shrubs, MANCH-000411!).
The species stands out by its small lobules with a
large, multicellular tooth, ca. 3–6 cells long and 2–4
cells wide, very thin-walled leaf cells with minute tri-
gones, no intermediate thickenings, and a smooth
cuticle, large bifid underleaves ca. 5× stem width and
with cuneate to rounded bases, and gynoecia on
elongate branches with 1(–2) innovations that remain
sterile or, occasionally, become fertile again (Gradstein
2021). The species is presumably dioicous; its androecia
are unknown. Lejeunea suffruticola keys out near L. flava
but the latter species is autoicous and usually copiously
fertile with androecia and gynoecia, the lobule tooth is
small, one-celled, and the leaf cells have distinct tri-
gones and intermediate thickenings, and a rough
cuticle. The record from the Galápagos Islands is the
second locality known of L. suffruticola and the first
from the Galapagos Islands. The Galápagos material
was previously published as L. setiloba Spruce (Grad-
stein 2009); the latter species differs from
L. suffruticola by the narrower lobule tooth (1–2 cells
wide), much smaller underleaves, 1.5–2.5× stem width,
and autoicous sexuality (Bastos and Gradstein 2020).
12. Lescuraea mutabilis (Brid.) Lindb. ex I.Hagen
Contributors. P. Saha, Md. N. Aziz and D. Maity
India. West Bengal: (1) Darjeeling, on way to Tiger Hill,
growing on rocks, in association with Hydrogonium
196 L. T. ELLIS ET AL.
amplexifolium (Mitt.) P.C.Chen and Ectropothecium ker-
stanii Dixon & Herzog, 27°00′168′′N, 88°15′862′′ E,
2347 m a.s.l., 03 December 2011, leg. Pamela Saha
64138c (CAL); (2) Jalapahar, growing on soil, in associ-
ation with Bellibarbula recurva (Griff.) R.H.Zander, 27°
01′50.4′′N, 88°15′58.5′′ E, 2231 m a.s.l., 18 July 2012,
leg. Pamela Saha 64184a (CAL).
The genus Lescuraea Schimp. is represented by nine
species worldwide (Crosby et al. 1999). Chopra (1975)
and Lal (2005) reported six species from India. In
India, the genus is well distributed in Western Hima-
laya. Vohra (1983) described one species (Lescuraea
darjeelingensis Vohra) from Darjeeling district.
Lescuraea mutabilis is characterised by: small plants,
forming dense tufts, slightly glossy, green; leaves lan-
ceolate with gradually long acuminate apex, margin
recurved faintly, dentate above; costa thick, percurrent;
median leaf cells linear; alar cells differentiated, few,
quadrate, thin-walled. The species L. mutabilis is
quite uncommon in India. It was previously reported
from Jammu and Kashmir in Western Himalaya.
Recently the species has been collected from Darjeel-
ing district of West Bengal. This is the first report of
L. mutabilis from Darjeeling district as well as from
the Eastern Himalaya.
13. Marsupella boeckii (Austin) Lindb.
Contributors. N. A. Konstantinova and A. N. Savchenko
Russia. Franz Josef Land Archipelago, Land Alexandra
Island, Severnaya Bay, remnant of the basalt plateau
(ca. 20 m high) with a large-column separation with
prismatic columns, 80°46′37′′N, 47°48′55′′ E, 22 m a.s.l.
In crevices and on ledges on soils and peat in cushions
among mosses and lichens. On edge of mats with
dominant Gymnomitrion corallioides Nees and admix-
ture of Trilophozia quinquedentata (Huds.) Bakalin,
Gymnomitrion concinnatum (Lightf.) Corda, Diplophyl-
lum albicans (L.) Dumort., 12 July 2019, leg. A. N. Sav-
chenko CA16-19-6a [KPABG(H): 123005, kpabg.ru›;cris/
?q=node/16]
The plants are quite characteristically red-brown,
with numerous stolons, small remote leaves, with a
single perianth. Cells of leaves with distinct trigones,
small ca. 12–18 μm.
This is a notable addition to the poorly known liver-
wort flora of the Franz Josef Land. The locality on Franz
Josef Land is situated ca. 60 km north of the previously
considered northernmost locality on Prins Oscar Land
(Nordaulandat, Svalbard) (Konstantinova and Sav-
chenko 2008).
14. Marsupella sparsifolia (Lindb.) Dumort.
Contributor. S. Ștefănuț
Romania. Southern Carpathians: (1) Ţarcu Mountains,
Mătania Peak, Hunedoara County, 45°18′55.8′′N, 22°
37′48.8′′E, 1774 m a.s.l., 16 September 2020, leg.
S. Ștefănuț,s.n.,det.S.Ștefănuț(BUCA B12085); (2)
45°18′53.3′′N, 22°37′44.4′′ E, 1796 m a.s.l., 16 Septem-
ber 2020, leg.S.Ștefănuț,s.n.,det.S.Ștefănuț(BUCA
B12086,B12094).
This is the second report of Marsupella sparsifolia for
Romania (Ellis, Asthana, et al. 2015). Marsupella sparsi-
folia grows on rocks in the eastern part of Mătania
Peak, along with other bryophytes such as Gymnomi-
trion concinnatum (Lightf.) Corda, Marsupella emargi-
nata (Ehrh.) Dumort. and Andreaea rupestris Hedw.
The nearest other locality for this species is in Bulgaria,
but this is data deficient (Hodgetts and Lockhart 2020).
The conservation status of M. sparsifolia in Romania is
changed from Critically Endangered –B1ab(ii,iii) + 2ab
(ii,iii) to Endangered –EN B2ab(ii,iii,iv).
15. Odontoschisma denudatum (Nees) Dumort.
Contributors. K. K. Rawat, V. Sahu and R. R. Paul
India. Uttarakhand, Rudraprayag, Tungnath near
temple, 30°29′20.23′′N, 79°13′01.24′′ E, 3465 m a.s.l.,
on soil, 18 June 2016, leg. K. K. Rawat s.n. (LWG
321601C).
Odontoschisma denudatum is reported for the first
time from Uttarakhand in the Western Himalaya.
Earlier this species was reported from Arunachal
Pradesh, Meghalaya and Sikkim in the Eastern Hima-
laya (Long 2005; Singh and Nath 2007; Rawat and
Verma 2014). The plants were yellowish to brown,
prostrate to erect, 1.2–1.6 mm long and 1–1.2 mm
wide including leaves, somewhat laterally compressed,
stolons frequent. Stem in cross-section rounded, 0.24–
0.28 mm in diameter, 10–12 cells high, epidermal cells
thick-walled, 30–40 in number, 20–32 µm long and 16–
20 µm wide, medullary cells larger than epidermal
cells. Leaves contiguous to imbricate, obliquely
inserted, ovate, 0.32–0.48 mm long and 0.40–
0.48 mm wide, apex rounded, margin entire. Leaf mar-
ginal cells isodiametric to hexagonal, 16–28 µm long,
20–40 µm wide, trigonous; median cells 20–28 µm
long, 20–32 µm wide, hexagonal, thick-walled, tri-
gones nodulose; basal cells 20–40 µm long, 20–
26 µm wide, rectangular to polygonal, trigonous.
Underleaves not seen.
16. Philonotis longiseta (Michx.) E.Britton
Contributor. E. Fuertes
Argentina. (1) Province Misiones, Depto Montecarlo,
de Montecarlo al Puerto de Paranay, bosque secun-
dario de la selva paranaense, 95 m a.s.l., 26°41′S, 54°
49′W, 25 September 2007, leg. E. Fuertes & C. Prada
s.n., det. E. Fuertes (MACB). (2) Province Jujuy,Departa-
mento Ledesma. P.N. Calilegua, paraje el Tigre, selva
montana con alisos (Alnus acuminata Kunth) las
Yungas, 1550–1790 m a.s.l., 1 June 2008, leg.
E. Fuertes & C. Prada s.n.det. E. Fuertes (MACB).
Plants green to yellowish-green, in loose tufts to
1.5–3 cm high, stem simple, ending with a whorl of
branches. Leaves straight or moderately secund,
JOURNAL OF BRYOLOGY 197
ovate or triangular lanceolate, 0.80–1.00(–1.40) × 0.18(–
0.25) mm; margin toothed, teeth double; costa percur-
rent or excurrent; upper cells rectangular 27–42(–45) ×
6–7.5 (–8.7) µm, papillose prorulate, median and basal
cells laxer, 33–55 × 10–12.5 µm; alar cells quadrate to
subrectangular 20–25 × 15–20 µm. Autoicous. Setae
1.5–2.0 (–2.30) cm long, capsule globose 2–2.5 (–3) mm
long; peristome double, exostome with 16 teeth
reddish, densely papillose above, finely papillose below,
endostome yellowish, papillose throughout. Spores sub-
reniform 25–27 × 20–22 µm, brownish, densely papillose.
Illustrations in Allen (1999:Figure3E–I)
In Argentina Philonotis longiseta was found on soils
in side banks, especially in moist and shaded forests. Its
range is Neotropical in two disjunct ecoregions: the
phytogeographic province Yungueña, located in NW
Argentina (Jujuy) and the phytogeographic province
Paranaense, located in the NE (Misiones), these consti-
tuting the two ecoregions with subtropical forests in
Argentina (Cabrera and Willink 1980). The global
range includes America, in three disjunct areas: North
America (USA, México); Central America (Guatemala,
Costa Rica); and N–NW South America, specifically
Venezuela (Griffin III 1994; Allen 1999), Colombia,
Ecuador Perú and Bolivia (Tropicos 2020) and NW–NE
Argentina (this contribution).
17. Polytrichum commune Hedw.
Contributor. H. M. H. van Melick
Chile. Capitan Prát Province in the Región Aysén (XI),
on the border of Nothofagus Blume forest on open,
sandy soil near a peat bog between Caleta Cachorro
and Lago Vergas, 30 m a.s.l., 47°40′24′′ S, 73°04′34′′ W,
20 January 2018, leg. H. M. H. van Melick 219327,
verified by Dr. F. Müller and Dr. J. Larrain.
The species is cosmopolitan. It is distinguishable
from all the other local species by the grooved terminal
cells of the lamellae. Of the many Polytrichum-like
specimens (which included Polytrichum juniperinum
Hedw., Polytrichum strictum Brid. and Polytrichadelphus
magellanicus (Hedw.) Mitt.) collected from several
locations in the fairly cool area, only one belonged
to Polytrichum commune. It is plausible that the
species is uncommon in Patagonia. The population
was sterile.
18. Pseudoparaphysanthus touwii (Ochyra & Enroth)
S.Olsson, Enroth, Huttunen & D.Quandt
Contributors. J. Enroth, J. R. Shevock and D. S. Salas
Philippines. Negros Occidental Province, Balinsasayao
Twin Lakes Natural Park, along Olayan Stream accessed
offan unimproved trail at the southern end of Balinsa-
sayao Lake then upslope toward Guinsayawan Peak, 9°
20′59.8′′N, 123°10′3.2′′ E, 1040 m a.s.l. Mixed hardwood
evergreen tropical rain forest along cascading stream
with volcanic bedrock, submerged in stream, rheo-
phyte, leg. James R. Shevock 56044 with Daryl S. Salas
(specimen examined H3241369; duplicates BRIT, CAS,
CMUH, E, H, KRAM, MO, NICH, NY, PNH, PTBG).
Pseudoparaphysanthus (Broth.) S.Olsson, Enroth,
Huttunen & D.Quandt (Neckeraceae) is a south-east
Asiatic genus of five rheophytic species previously
placed in Neckeropsis Reichardt sect. Pseudoparaphy-
santhus (Broth.) M.Fleisch. (Ochyra and Enroth 1989;
Olsson et al. 2016). The genus is morphologically
characterised by rigid plants with a very strong costa
in the leaves, and often a strong, bi- to multistratose
limbidium along the leaf margins. These characters
are adaptations to harsh rheophytic habitats (Enroth
1999; Shevock et al. 2017).
Pseudoparaphysanthus touwii was previously
known from two specimens from Papua New Guinea
(Ochyra and Enroth 1989; Enroth 1992). Its most dis-
tinctive characters are a 2–3 stratose lamina and 4–8
stratose marginal limbidia. The Philippine specimen
has these characters, although in some other respects
it differs from the New Guinean specimens. The leaves
in the New Guinean material are mostly oblong-ovate
with broadly acute to obtuse apices, and the upper
laminal cells are to 20 µm long, becoming to 30 µm
at midleaf and to 50 µm long towards the leaf base.
The Philippine specimen has mostly lanceolate or
oblong-lanceolate leaves with more acute apices,
the upper laminal cells are to 30 µm long, at midleaf
to 50 µm and near leaf base to 80 µm long. Rheophy-
tic mosses are known to be very variable, and since
the basic leaf structure in the New Guinean and Phi-
lippine specimens is similar and only known from
P. touwii, we are satisfied that they represent the
same species.
19. Ptychomitrium incurvum (Schwägr.) Spruce
Contributors. S. Poponessi and A. Cogoni
Sardinia. Tavolara Island, Province of Sassari, a Site of
Community Importance (SCI) ITB010010 Isola di Tavo-
lara, Molara e Molarotto, 40°8.931′N, 9°68.688′E, ca.
195 m a.s.l., on soil, 20 June 2004, leg. A. Cogoni, det.
S. Poponessi (CAG SA4.5.1.1).
The record derives from a project studying the
smaller islands offSardinia to monitor climate change
and how this might affect bryophyte diversity. Ptycho-
mitrium incurvum is a submeridional/mountain-temper-
ate species narrowly distributed in Europe (Dierßen
2001). It is considered Critically Endangered (CR) -
B2ab (ii, iv, v) in Europe and Data Deficient (DD) in
Italy according to Hodgetts and Lockhart (2020). This
is the first report from Italy in the last 70 years; the pres-
ence in Piedmont and Lombardy is based only on old
records according to Aleffiet al. (2020). Ptychomitrium
incurvum is considered Regionally Extinct in Spain
(Casas et al. 2009; Brugues et al. 2014) and was recently
observed in France in the Pyrenees-Orientales (Hugon-
not and Chavoutier 2016). It is widespread in eastern
North America, from Canada (southern Ontario) south
198 L. T. ELLIS ET AL.
to the Gulf States and Texas and west to Iowa and
Kansas (Crum and Anderson 1980).
The material collected was deep black, in small,
rounded cushions growing under Cistus monspeliensis
L., the largest tufts reaching 0.4–0.6 cm high, with
strongly crisped and obtuse-entire leaves. The leaves
were characteristically concave at the apex and cucul-
late. Specialised asexual reproduction was rare with
short-uniseriate or branched gemmae, several cells
long, on branched axillary filaments.
20. Ptychostomum marratii (Hook. & Wilson)
J.R.Spence
Contributors. A. Uygur, T. Ezer, S. Karaman Erkul and
M. Alataş
Turkey. Mersin Province, Taşeli Plateau, BozyazıDis-
trict, Akcami Village: 36°9′N, 32°54′E, growing on cal-
careous muddy soil, 130 m a.s.l., 16 August 2020, leg.
and det. T. Ezer, A. Uygur & M. Alataşs.n. (A.U.1800).
Hitherto, Ptychostomum Hornsch. has been rep-
resented in Turkey by 17 taxa (Kürschner and Frey
2020a), the number increasing to 18 with this new
record. Known in Europe as Baltic Bryum, P. marratii
is identified by its blunt-tipped, strongly concave,
oval-oblong leaves and obscurely bordered leaf
margin; costa ending below apex; basal leaf cells rec-
tangular or slightly rhomboid, and bases of the
leaves not reddish. Ptychostomum marratii is similar
to P. calophyllum (R.Br.) J.R.Spence. However,
P. calophyllum prefers different habitats and has
broader and less concave leaves (Smith 2004).
The study area is characterised by a Mediterranean
climate with mild wet winters and warm to hot, dry
summers. The population of Ptychostomum marratii
was sterile and sporophytes are unknown in Turkey.
Ptychostomum marratii is circumpolar, subneutro-
phytic, hygrophytic and photophytic. Its preferred
habitat is slightly saline, on damp, muddy soil in dune
slacks and by the sea (Dierßen 2001;Smith2004). The
present specimens were collected on calcareous
muddy soil next to a small stream. Although the
species is found in Europe (Britain, Denmark, Latvia,
Estonia, Finland, Norway, Faroes, The Netherlands,
France, Germany, Iceland, Ireland, Slovakia, Sweden), it
is included in the IUCN Red List of Threatened Species
in the EN category (Hodgetts, Blockeel, et al. 2019).
Outside Europe, it is reported only from north-central
Asia (the Altai Mountains) and North America (Smith
2004;Kekes2009;Šoltés et al. 2010;Rosetal.2013).
In addition, P. marratii is known from Yemen and Syria
(Callaghan and Farr 2018; Kürschner and Frey 2020b).
21. Pyramidula tetragona (Brid.) Brid.
Contributors. M. Kropik and H. G. Zechmeister
Austria. Bohemian Massif, steep slope at the river
Thaya east of Hardegg, 48°51′11′′N, 15°52′33′′ E(WGS
84), 365 m a.s.l, two cushions with sporophytes and
calyptrae on open clayey soil in a dry grassland
around rock outcrops, 8 April 2020, leg. &det.: Michaela
Kropik and Harald Zechmeister s.n. (Private herbarium
of Harald Zechmeister, University of Vienna). Associ-
ated bryoflora: Abietinella abietina (Hedw.) M.Fleisch.
var. abietina;Didymodon acutus (Brid.) K.Saito;
Entosthodon fascicularis (Hedw.) Müll.Hal.; Microbryum
curvicollum (Hedw.) R.H.Zander; Phascum cuspidatum
Hedw. var. cuspidatum;P. cuspidatum var. piliferum
(Hedw.) Hook. & Taylor; Riccia sorocarpa Bisch.;
Weissia controversa Hedw.
Pyramidula tetragona is at risk of extinction in
several countries and regions. Intensification of agri-
culture, intensive use of herbicides and fertilisers and
encroachment of shrubs on abandoned grasslands
have decreased the extent of suitable habitats and
resulted in small, isolated subpopulations (Campisi
and Cogoni 2019). Long-distance dispersal and recolo-
nisation of new habitat are extremely limited for P.tet-
ragona due to its large spores and asexual
underground tubers (Høitomt et al. 2012).
In the European Red List of mosses, Pyramidula
tetragona has been assessed as EN (Campisi and
Cogoni 2019). Most countries neighbouring Austria
report small and endangered populations: see
Papp et al. (2000,2010)forHungary;Kučera et al.
(2012)andDřevojan et al. (2019)fortheCzech
Republic; Mišíková et al. (2019)forSlovakia;
Caspari et al. (2018)forGermany;andAleffiet al.
(2008) for Italy. No recent records are listed for Swit-
zerland (Schnyder et al. 2004; Swissbryophytes 2020)
or for Slovenia (Martinčič2016). In Austria, Juratzka
(1882) reported the last observations of P.tetragona
in Lower Austria close to Sankt Pölten, in Styria at
Liebenau close to Graz, and in the Tyrol close to
Lienz. Wallnöfer (1888) added a further site in Car-
inthia close to Klagenfurt; Dalla Torre and Sarnthein
(1904) in the Tyrol at Innsbruck. None of these
reports has been reconfirmed since then. Therefore,
this is the first record of P. tetragona in Austria for
more than 100 years.
22. Rhytidiadelphus loreus (Hedw.) Warnst.
Contributor. Cs. Németh
Hungary. Somogy County, [9871.3] (Central European
Mapping Scheme), BelsőSomogy, ca. 3.4 km south of
the village Rinyabesenyő, on acidic soil at the edge
of oak forest, ca. 155 m a.s.l., 46°08′11.6′′ N, 17°
30′57.0′′E, associated with Ceratodon purpureus
(Hedw.) Brid., Polytrichum formosum Hedw., Polytri-
chum piliferum Hedw., 29 November 2020, leg. &det.
Cs. Németh s.n., conf. P. Erzberger (Herb. Cs. Németh
10276).
Rhytidiadelphus loreus is an acidophilic species
withapredominantlyasuboceanic-montanedistri-
bution in Europe. It has been found in all the
countries surrounding Hungary (Hodgetts and
JOURNAL OF BRYOLOGY 199
Lockhart 2020). Rhytidiadelphus loreus is missing
from the latest checklist of Hungarian bryophytes
(Papp et al. 2010).
23. Riccia fluitans L.
Contributors. M. Puglisi and S. Sciandrello
Sicily. Torrente Vacco Minore (Fiumedinisi, Peloritani
Mts.), 38°03′26.88′′N, 15°21′21.33′′ E, 529 m a.s.l., on
wet and shady rocky outcrops facing north together
with Oxyrrhynchium schleicheri (R.Hedw.) Röll., Conoce-
phalum conicum (L.) Dumort., 16 November 2019, leg.
S. Sciandrello s.n., det. M. Puglisi (CAT).
Riccia fluitans is one of the most common species of
Riccia L. and has a worldwide distribution; it is known
from Europe, Asia, northern America, northern Africa
and New Zealand. Nevertheless, despite its wide distri-
butional range, in Europe it is considered endangered
in many countries (Hodgetts and Lockhart 2020). In
Italy, it is quite well distributed, but hitherto has not
been found in Sicily. In the new locality it occurred
on rocky outcrops by water in the Peloritani Moun-
tains, confirming the importance of the Mediterranean
mountains for species conservation (e.g. Puglisi et al.
2012,2013,2014,2015; Sciandrello et al. 2015,2019;
Puglisi and Privitera 2018). The new locality is charac-
terised by a chasmophytic vegetation, rich in bryo-
phytes and pteridophytes associated with siliceous,
dripping cliffs, mainly dominated by Osmunda regalis
L., Athyrium filix-femina (L.) Roth, and Adiantum capil-
lus-veneris L., referable to Adianto capilli-veneris-
Osmundetum regalis Brullo, Lo Guidice & Privitera
1989 (Adiantetea capilli-veneris Br.-Bl. in Br.-Bl., Rous-
sine & Négre 1952).
24. Riccia lamellosa Raddi
Contributor. E. A. Borovichev
Russian Federation. Western Siberia, Republic of
Altai, Saylyugemsky National Park, Kosh-Agachsky Dis-
trict, Kosh-Agach Settelment, 49.975110°N, 88.671558°
E, steppe vegetation with rock fields, under stone, 22
May 2019, leg. E. Borovichev #RA 6-20 (INEP).
This is the first record of Riccia lamellosa for the
Republic of Altai (cf. Konstantinova et al. 2009). The
species has a very wide distribution, and is known in
Mediterranean Europe, North and Central Africa and
the southern part of North America, as well as temper-
ate and subtropical parts of South America and Asia
(Ladyzhenskaja 1961; Jovet-Ast 1986; Söderström
et al. 2002; Katagiri and Furuki 2012; Borovichev and
Bakalin 2016;Parolly et al. 2018; Özenoğlu Kiremit
et al. 2019). According to Gradstein (2017)Riccia lamel-
losa is characterised by an almost perfect circum-
amphi-tropical range, occurring disjunctively through-
out the Mediterranean and subtropical regions of the
world. The records of Riccia lamellosa nearest to the
Altai locality are in the Gissarskiy Range (Tajikistan;
Zerov et al. 1974), Akmolinsk Region (Kazakhstan;
Ladyzhenskaja 1961) and the Republic of Buryatiya
(Russia; Bakalin and Taran 2004).
25. Riccia subbifurca Warnst. ex Croz.
Contributors. E. A. Borovichev, N. R. Shafigullina
Russian Federation. Central part of European Russia,
Republic of Tatarstan, Zelenodol’skiy District, 55°
52′19′′N, 48°40′34′′ E, Aysha Settlement vicinity, along
highway #A295, meadow edge of the arable field, on
soil among grass, 10 November 2017, leg.
N. Shafigullina #14 (INEP(H)-201).
This is the first report of Riccia subbifurca for Russia
(cf. Konstantinova et al. 2009). Most thalli were fertile
and occurred in incomplete rosettes. The species has
a mainly Mediterranean and sub-Mediterranean distri-
bution (Jovet-Ast 1986; Dierßen 2001; Özenoğlu
Kiremit et al. 2019). It is recorded from many localities
in Europe, in Africa from Algeria and Morocco and in
Asia from Iran, Israel, Syria, Turkey and the United
Arab Emirates (Schumacker and Váňa2005; Özenoğlu
Kiremit 2011; Hodgetts 2015; Özenoğlu Kiremit et al.
2019). In the countries of the former USSR it is
known only from Georgia (Jovet-Ast 1986; Parolly
et al. 2018).
26. Riella affinis Howe & Underw.
Contributor. J. G. Segarra-Moragues
Niger. Air Mountains, Tamgak, Agoum, 19°01′N,
8°33′E, Guelta, en mélange avec Characées, algues
filamenteuses et Diatomées, Mars 1977, leg. Henri
Dumont 143E, 165261(BR-BRYO).
Recent studies on the genus Riella Mont. have
revealed some new species and populations of
poorly known species worldwide (Puche and
Segarra-Moragues 2013; Segarra-Moragues and
Puche 2014; Segarra-Moragues et al. 2014,2019a,
2019b). Difficulties in addressing taxonomical and bio-
geographical studies in Riella arise from the scarcity of
collections which are often scattered across herbaria.
The Biodiversity Information Facility (GBIF) may help
locating specimens across herbaria. A GBIF search
revealed a Niger specimen of Riella cossoniana Trab.
of unusual ecology (Compère 1980) and separated
from other R. cossoniana records. Examination of the
specimen revealed it matched the monoicous
R. affinis. In monoicous Riella, the antheridia are
scarce, produced in small groups rather than in con-
tinuous series. Riella affinis is protandrous, with
antheridia located in the sinuses of the thallus wing
(Segarra-Moragues et al. 2014). The Niger voucher
consisted of small plants in the male phase and one
fully fertile plant, with antheridia and winged female
involucres. It seems that antheridia in this plant
passed unnoticed, and thus it was interpreted as a
female of the dioicous R. cossoniana. Such misidentifi-
cations have been reported in this species (Segarra-
Moragues et al. 2019a).
200 L. T. ELLIS ET AL.
Riella affinis extends from the Canary Islands to India
and a single South African population. Its distribution
is highly scattered with only 10 populations known
(Segarra-Moragues et al. 2019a). Unlike the halophilous
R. cossoniana,R. affinis develops submerged in fresh-
water ponds and lakes. The Niger population occurred
in a guelta, a small ephemeral rock pool of Saharan
ravines, as did another Algerian population (Puche
and Boisset 2009).
27. Sarmentypnum trichophyllum (Warnst.) Hedenäs
Contributors. G. Brusa, G. Bergamo Decarli,
F. Sguazzin, M. Aleffiand S. Poponessi
Italy. (1) Lombardy Region: Ballone Lake (Val di Lei),
Province of Sondrio. 46°24′33′′N, 9°25′16′′ E, ca. 2320
m a.s.l., submerged on the shores of the lake, 17 July
1996, leg. G. Brusa s.n. (priv. herb.); (2) Trentino-Alto
Adige Region: Malga Binasia (Bresimo) Province of
Trento, 46°26′38′′N, 10°56′46′′ E, floating in the Malga
Binasia spring, 2225 m a.s.l., September 2017, leg.
G. Bergamo Decarli, det. F. Sguazzin (priv. herb.); (3)
Passo del Rombo (Moso in Passiria) Province of
Bolzano, 46°53′34′′N, 11°05′51′′ E, submerged in small
lakes between 2547 and 2575 m a.s.l., September
2018, leg. G. Bergamo Decarli, det. F. Sguazzin (priv.
herb.); (4) Marschnell Moor (Ultimo) Province of
Bolzano, 46°34′21′′N, 10°58′26′′ E, submerged in the
marsh, 2345 m a.s.l., October 2018, leg. G. Bergamo
Decarli, det. F. Sguazzin (priv. herb.); (5) Ochsenberg
Alm (Curon Venosta) Province of Bolzano, 46°47′46′′N,
10°36′28′′E, submerged in the lake, 2154 m a.s.l.,
October 2018, leg. G. Bergamo Decarli, det. F. Sguazzin
(priv. herb.); (6) Westklapfberg Tal (Ultimo) Province of
Bolzano, 46°27′35′′N, 10°54′19′′E, submerged in the
lake, 1935 m a.s.l., October 2018, leg. G. Bergamo
Decarli, det. F. Sguazzin (priv. herb.).
Hedenäs (2006) transferred this dioicous species of
WarnstorfiaLoeske to Sarmentypnum Tuom. &
T.J.Kop. based on the results of that paper and those
previously published by Hedenäs et al. (2005). In the
Mediterranean area (including Italy) this genus is rep-
resented to date by Sarmentypnum exannulatum
(Schimp.) Hedenäs and S. sarmentosum (Wahlenb.)
Tuom. & T.J.Kop.During a series of research studies
conducted in the central-eastern Alps, Sarmentypnum
trichophyllum was found in Lombardy and in some
localities of Trentino-Alto Adige. The specimens col-
lected grew mainly submerged or floating in small
alpine lakes, in mineral-rich water.
The species occurs in many localities in Sweden,
Finland and Norway, in the Svalbard Islands, Russian
Federation, Latvia; Canada (Alaska), United States of
America, Greenland; Ecuador, Colombia. The only
report for central and southern Europe is in Austria
(Tirol, Ötztaler Alp bei Schoenwieshuette unter der
Rotmoosferner, Pitztal, Tallage, leg. A. Blaufuss,
01.08.1972) (Grims 1999). Therefore, this is the first
report of Sarmentypnum trichophyllum for Italy and
the second for central and southern Europe. The
present report of S. trichophyllum in Italy is an impor-
tant confirmation and extension of range southwards.
In addition, the species is included in the IUCN Red List
in the LC category (Hodgetts, Calix, et al. 2019)
28. Scapania crassiretis Bryhn
Contributor. S. Ștefănuț
Romania. Southern Carpathians: FăgărașMountains,
Viștișoara glaciar ring, Brașov County, 45°36′49.1′′N,
24°45′48.6′′E, 2070 m a.s.l., 20 August 2020, leg.
S. Ștefănuț,s.n.,det.S.Ștefănuț(BUCA B12023,B12028).
This is the first report of Scapania crassiretis from
Romania in the last 120 years (Ștefănuț2008;Ștefănuț
and Goia 2012;Ștefănuț2020). The specimens of
S. crassiretis from FăgărașMountains, Negoiu Moun-
tain, 2200 m a.s.l., leg. &det. K. Loitlesberger, 16
August 1897, conf.S.Ștefănuț, 20 October 2014 (W
1898-8258) was the previous single report of this
species for Romania (Loitlesberger 1898;Ștefănuț
2020).
Scapania crassiretis grew along with other bryo-
phytes such as Fuscocephaloziopsis albescens (Hook.)
Váňa & L.Söderstr. var. albescens,Anoectangium aesti-
vum (Hedw.) Mitt., Distichium inclinatum (Hedw.)
Bruch & Schimp.,Flexitrichum gracile (Mitt.) Ignatov &
Fedosov, Meesia uliginosa Hedw. and Myurella tener-
rima (Brid.) Lindb. The nearest other locality of this
species is in Bulgaria (Hodgetts and Lockhart 2020).
The conservation status of S. crassiretis in Romania is
Critically Endangered –CR B2ab(ii,iii).
29. Scapania kaurinii Ryan
Contributor. M. V. Dulin
Russian Federation. Komi Republic, subpolar Urals,
Intinsky District, ‘Yugyd Va’National Park, near the
Bolshoi Balbanty Lake and the ‘Zhelannaya’tourist
camp, top of the Barkova Mountain, 65°11′00.1′′N,
60°16′25.2′′E, ca. 1217 m a.s.l., rock outcrops near
groundwater outlet, on wet ground between stones,
among other liverworts, Gymnomitrion concinnatum
(Lightf.) Corda, Lophozia murmanica Kaal., Lophozia ven-
tricosa (Dicks.) Dumort., Ptilidium ciliare (L.) Hampe, Sca-
pania spitsbergensis (Lindb.) Müll.Frib., Tetralophozia
setiformis (Ehrh.) Schljakov, and Trilophozia quinqueden-
tata (Huds.) Bakalin, some plants with gemmae, anther-
idia, perianths, 9 July 2009, leg. &det. M. V. Dulin 819mvd
(58150, 58151, 58152, 58143) (SYKO).
This is the first report of Scapania kaurinii from the
Komi Republic. It is an arctomontane species with an
uncertain distribution (Konstantinova 2000).This liver-
wort is assessed as Vulnerable in both Europe and the
EU 28 and included in the Red Data Book of some
regions: Finland, Norway and the Russian Federation
(Murmansk Province) (Konstantinova 2014; Halling-
bäck et al. 2019).
JOURNAL OF BRYOLOGY 201
Scapania kaurinii is known from several countries in
Europe (Finland, Norway, Sweden) and from some
localities in Svalbard and Jan Mayen. It is also reported
from North America (Alaska, Ellesmere Island) and
Greenland (Damsholt 2002; Hallingbäck et al. 2019),
and according to the GBIF database, is known in
Canada (British Columbia, Quebec). In Russia, it is
recorded in Murmansk Province (Khibiny Mountains,
Nyavka-Tundra Mountains), Arhangelsk Province
(Novaya Zemlya Island), Yamal-Nenets Autonomous
District (Yamal Peninsula), Khanty-Mansi Autonomous
District (Ner-Oika Mountain), Sverdlovsk Province
(northern Urals), Krasnoyarsk Krai (Taymyr Peninsula),
Irkutsk Province (Udokan Range), Buryatia Republic
(Topografov Peak), Republic of Sakha (Yakutia)
(Yudomo-Mayskoe Upland, Suntar-Khayata, Tokinsky
Stanovik, Tas-Kystabyt Ranges), Zabaikal’sky Krai
(Kodar Range), Magadan Province (Kolyma Upland),
Chukchi Autonomous District (western and eastern
Chukotka, Anadyr River basin), Kamchatka Krai (north
and central Kamchatka) and Sakhalin Province (Nabils-
kij Range). Records of Scapania kaurinii from the Kam-
chatka Peninsula, Novaya Zemlya and the Yamal
Peninsula are based on identifications of sterile collec-
tions and remain dubious (Bakalin 2004; Konstantinova
et al. 2009; Choi et al. 2012; Konstantinova 2014; Kon-
stantinova and Lapshina 2014; Sofronova 2011,2013,
2018,2019; Sofronova et al. 2013,2015; Sofronova
and Potemkin 2016).
30. Scapania obscura (Mitt.) Müll.Frib.
Contributors. A. D. Potemkin, M. Ryan and
T. T. McIntosh
Canada. Yukon, Richardson Mountains, 67.777738°N,
136.690401°E, ca. 800 m a.s.l., оn soil over rock below
waterfall, 29 July 2019, leg. T. McIntosh, det.
A. D. Potemkin and M. Ryan (LE, UBC).
Scapania obscura is a rare species of the section Sca-
pania (Dumort.) Dumort. It has been previously
reported in North America from Canada: British Colum-
bia, Northwest Territories, and Nunavut, and from the
US: Alaska, Oregon, and Washington (Potemkin 2016).
The record from Yukon fills a gap in the distribution
of this species between Alaska and Northwest Terri-
tories. The species is exceedingly malleable and devel-
ops various morphotypes (Potemkin 1998,pp.37–38,
46, Figures 3–5) that may result in misidentifications.
31. Scapania scandica (Arnell & H.Buch) Macvicar
Contributors. A. D. Potemkin and M. Ryan
Canada. Yukon, Richardson Mountains, 67°31′16.4′′N,
136°30′18.1′′E, 788 m a.s.l., on humus on side of
opening of rodent den amongst rocky talus in
tundra, 29 July 2019, leg. M. Ryan s.n., det.
A. D. Potemkin and M. Ryan (LE, UBC).
Scapania scandica is an infrequent and quite vari-
able species of the section Curtae (Müll.Frib.) H.Buch,
rather widespread in northern North America, but pre-
viously not reported from Yukon (Potemkin 2016). The
record of this species from Yukon fills the gap in its dis-
tribution between Alaska and the Northwest
Territories.
32. Scapania spitsbergensis (Lindb.) Müll.Frib.
Contributor. M. V. Dulin
Russian Federation. Komi Republic, subpolar Urals,
Intinsky District, ‘Yugyd Va’National Park, near the
Bolshoi Balbanty Lake and the ‘Zhelannaya’tourist
camp, top of the Barkova Mountain, 65°11′00.1′′N,
60°16′25.2′′E, ca. 1217 m a.s.l., rock outcrops near
groundwater outlet, on wet ground between stones,
among other liverworts: Gymnomitrion concinnatum
(Lightf.) Corda, Lophozia murmanica Kaal., Ptilidium
ciliare (L.) Hampe, Scapania kaurinii Ryan, Trilophozia
quinquedentata (Huds.) Bakalin, 9 July 2009, leg. &
det. M. V. Dulin 819mvd (SYKO 58152).
This is the second report of Scapania spitsbergensis
from the Komi Republic. Earlier it was found in the
northern Urals (near the Halmersale Mountain) (Sofro-
nova et al. 2016). It is an arctomontane species with a
subcircumpolar distribution (Konstantinova 2000) and
was assessed as Vulnerable (VU) in Europe and in the
EU 28 as Endangered (EN) (Hallingbäck 2019). It is
included in the Red Data Book of Murmansk Province
and Komi Republic (Russia) (Konstantinova 2014;
Degteva 2019).
Scapania spitsbergensis is known from several
countries in Europe (northern Norway, Finland,
Sweden) and from some localities in Svalbard and
Jan Mayen. It is also reported from North America
(Alaska, Northwest Territories, Yukon, Ontario and
Maine) and Greenland (Damsholt 2002; Klimova and
Bakalin 2017; Hallingbäck et al. 2019). In Asia it is
known in Mongolia (Tsagaan Chuluut Mountain)
(Tsegmed et al. 2012).
In Russia, it is recorded in Murmansk Province,
Arhangelsk Province, Khanty-Mansi Autonomous Dis-
trict, Altai Republic, Krasnoyarsk Krai, Irkutsk Province,
Buryatia Republic, Republic of Sakha (Yakutia), Zabai-
kal’sky Krai, Amur Province, Chukchi Autonomous Dis-
trict, Magadan Province, Khabarovsk Krai and
Kamchatka Krai (Damsholt 2002; Konstantinova et al.
2009; Choi et al. 2012; Tsegmed et al. 2012; Sofronova
et al. 2013,2015; Konstantinova 2014; Konstantinova
and Lapshina 2014; Fedosov et al. 2015; Sofronova
2015,2018,2019; Bakalin et al. 2016; Sofronova and
Potemkin 2016; Klimova and Bakalin 2017)
33. Schistidium dupretii (Thér.) W.A.Weber
Contributors. S. V. O’Leary and P. Martin
Andorra. Bony de Les Neres, Ordino. 42°32′49.0′′N, 1°
33′49.2′′E, ca. 2210 m a.s.l., on concrete base of trans-
mitter station near mountain peak, 16 May 2019, leg.
and det. S. V. O’Leary & P. Martin s.n., conf. T. L. Blockeel.
202 L. T. ELLIS ET AL.
A small member of the genus, which can be con-
fused in the field with Schistidium confertum (Funck)
Bruch & Schimp.,from which it differs in its longer
capsule and leaves with very short hair-points. It is
rare in the Iberian Peninsula (Brugues and Guerra
2015), and a montane species throughout its range
in Europe (Blockeel et al. 2014). Blom (1996) notes
that, as a pioneer species of calcareous rock, it is very
common in man-made habitats, ‘on concrete by road-
sides, foundations of electric pylons’. In Bony de Les
Neres the specimens were collected from just such a
habitat.
34. Schistochilopsis grandiretis (Lindb. ex Kaal.)
Konstant.
Contributor. V. Hugonnot
Italy. Trentino-Alto Adige, Belluno, Lago di Sorapis,
northern slope, 300 m south of the lake 2000–2050 m
a.s.l., 46°31′09.3′′N, 12°13′24.1′′E, 28 June 2019, leg.
V. Hugonnot s.n.(Herb.VincentHugonnot).
For Italy, Schistochilopsis grandiretis was historically
reported in 1896 in Trentino-Alto Adige (Bisang
1991), and not seen since (Aleffiet al. 2020). A rare
arctic-alpine, circumpolar species, it has a sporadic
occurrence in Europe, mostly in Scandinavia and the
Alps.
Often associated with tundra in northern regions, in
Italy, it was observed in the alpine belt between dwarf
shrubs and herbs (Dryas octopetala L., Rhodothamnus
chamaecistus (L.) Rchb. etc.) on a calcareous slope on
humic, basic dolomitic soil with a lengthy snow
cover, where cryonival phenomena are predominant.
Associated bryophyte species included Distichium
capillaceum (Hedw.) Bruch & Schimp., Ditrichum
gracile (Schwägr.) Hampe, Blepharostoma trichophyl-
lum subsp. brevirete (Bryhn & Kaal.) R.M.Schust.,
Mnium thomsonii Schimp. and Pohlia cruda (Hedw.)
Lindb., which can be referred to the Distichion capilla-
cei vegetation type. Schistochilopsis grandiretis was
fully fertile there and seemed to benefit from natural
disturbance of the ground. This type of vegetation
has a permanent character and is of high conservation
value.
Schistochilopsis grandiretis is characterised by its
pure green colour, large midleaf cells (often more
than 60 µm wide), and a distinct purple strip running
along the ventral side of the stem. This deep colour-
ation extends to the base of the leaves or the apex
of the plant. It is distinct from the other two green
species of the genus known in the Alps, Schistochilopsis
incisa (Schrad.) Konstant. and S. opacifolia (Meyl.) Kon-
stant., which are also acidophiles that mostly do not
attain such high altitudes. Schistochilopsis grandiretis
has been reported from several low-altitude localities
in Germany and Belgium (Schumacker 1993) but the
taxonomic status of these populations should be re-
evaluated.
35. Schwetschkea usambarica Broth.
Contributor. J. Enroth
Uganda. Kibale, Oate’s Track, trail side in partial shade,
on Acanthus L. trunk, 1500 m a.s.l., 25 November 1983,
leg. Matti Nummelin s.n. (H3241370).
Buck and Pôrto (2010) revised the African species of
Schwetschkea Müll.Hal. and placed the genus in the
Brachytheciaceae. Three species were recognised:
S. fabronioides (Welw. & Duby) Broth., S. grateloupii
(Mont.) Müll.Hal. and S. usambarica Broth. The last-
mentioned can be distinguished from the other two
by several characters, such as the clearly shorter
upper laminal cells and the sparsely hairy vs. naked
calyptrae. The specimen reported here is a good one,
with both mature and young sporophytes. Some of
the young capsules still have the calyptrae attached,
and the lower calyptra portions bear long hairs, as illus-
trated by Buck and Pôrto (2010).
Schwetschkea usambarica is known from Kenya,
Malawi, Tanzania, Democratic Republic of the Congo,
and Zambia (O’Shea 2006). It is thus restricted to
eastern Central Africa. Buck and Pôrto (2010) regarded
is as ‘rare’and indeed only few collections are known.
The type from Tanzania is in V. F. Brotherus’herbarium
in H (https://plants.jstor.org/stable/10.5555/al.ap.
specimen.h3301270).
36. Southbya gollanii Steph.
Contributors. K. K. Rawat, V. Sahu and R. R. Paul
India. Arunachal Pradesh, Tawang, on soil, 12 Novem-
ber 2019, leg. K. K. Rawat and R. R. Paul s.n. (LWG
325635A).
Southbya gollanii [≡Gongylanthus gollanii (Steph.)
Grolle] is reported for the first time from state of Aruna-
chal Pradesh, in the Indian Himalaya. It has previously
been reported from Meghalaya and Uttarakhand (Ste-
phani 1906; Kashyap and Chopra 1932;Chopra1943;
Parihar 1961–1962; Kachroo 1970;Grolle1986;Long
and Grolle 1990; Parihar et al. 1994;Mizutanietal.
1995; Bapna and Kachroo 2000; Srivastava and Verma
2005; Singh et al. 2016). The plants are yellowish, 3–6
mm long, 1.0–1.5 mm wide including leaves, prostrate;
rhizoids numerous, hyaline, present on ventral surface.
Leaves closely imbricate, broadly ovate with rounded
apex, entire margins, 1.3–1.6 mm long, 1.1–1.6 mm
wide, margin recurved at apex; marginal cells 60–100
µm long, 24–36 µm wide, polygonal to rectangular,
thin-walled; median cells hexagonal, 60–112 µm long,
40–60 µm wide, thin-walled with small trigones; basal
cells 60–120 µm long, 40–60 µm wide, hexagonal;
cuticle smooth. Underleaves absent. Gemmae absent.
Sporophytes not found.
37. Sphagnum compactum Lam. & DC.
Contributors. E. Yu. Kuzmina and N. S. Liksakova
Russia. Sakhalin Region, Urup Island, southern part of
the island: (1) 6.03 km east of the mouth of the Kama
JOURNAL OF BRYOLOGY 203
River, 45.6519402°N, 149.5459702°E, 248 m a.s.l., peat-
land with ridge-pool complexes, hollow near the lake,
as admixture to Sphagnum fallax (H.Klingrr.) H.Klinggr.,
4 September 2019, leg. N. Liksakova 20, det. E. Kuzmina
(LE); (2) 6.23 km east of the mouth of the Kama River,
45.64912954°N, 149.5490847°E, 241 m a.s.l., peatland
with ridge-pool complexes, narrow ridge between
lakes, sedge-Sphagnum community dominated by
Carex middendorffiiF.Schmidt with an admixture of
tundra species, sometimes abundant, 4 September
2019, leg. N. Liksakova 23, det. E. Kuzmina (LE).
Sphagnum compactum is widespread in the Arctic
and Boreal Zones of the Holarctic. It is a rare species
for the Southern Kuril Islands, previously known only
from Iturup Island (Bakalin et al. 2009).
38. Sphagnum cuspidatum Ehrh. ex Hoffm.
Contributors. M. Kırmacı, F. Filiz and U. Çatak
Turkey. Province Giresun, Kürtün, between Kızılali
Yayla and İnçayırı, 40°49′15.4′′N, 39°02′33.9′′ E, 1650 m
a.s.l., in high mountain flushes, 31 August 2016, leg.
M. Kırmacı, F Filiz & U Çatak (MKIR 7532) (AYDN).
This new record raises to 25 the number of Sphag-
num L. taxa recorded for Turkey. All collections were
made in the northern part of the country, which is
influenced by an oceanic climate with a high annual
rainfall. There are limited peatlands over 1500 m a.s.l.
in the eastern Black Sea region (Kırmacıet al. 2019),
these areas being among the more fragile ecosystems
of Turkey and potentially affected by climate change.
Sphagnum cuspidatum is found mostly in ombro-
trophic bogs or acid fens but may also grow in
ditches or oligotrophic flushes (Daniels and Eddy
1985). The Turkish specimen was collected from high
mountain flushes among vegetation composed of
mixed forests consisting of Picea orientalis (L.) Link.
and Fagus orientalis Lipsky. Rhododendron ponticum
L. and Vaccinium L. sp. are common in the lower layer
and in open forest areas. Various hydrophytic herbs
and Asplenium L. sp. are common around flushes.
These taxa are known to develop on acidic soils. Sphag-
num centrale C.E.O.Jensen and Polytrichum commune
Hedw. are the other dominant associated bryophytes
in the area. As with other aquatic taxa, Sphagnum cuspi-
datum is highly variable. Aquatic forms, however, can
easily be separated by their fine, delicate morphology
(Daniels and Eddy 1985). Our specimen is closer to a ter-
restrial form, which is relatively difficult to distinguish
and can easily be overlooked among other taxa. The
characteristic feature of the terrestrial form is the rela-
tively long stem and side branch leaves and the
tightly curled ends of the cuspidate side branch leaves.
39. Sphagnum pulchrum (Lindb. ex Braithw.) Warnst.
Contributors. E. Yu. Kuzmina and N. S. Liksakova
Russia. Sakhalin Region, Urup Island, southern part of
the island, 5.94 km east of the mouth of the Kama
River, 45.65420587°N, 149.5444681°E, 253 m a.s.l.,
edge of the peatland with ridge-pool complexes,
sedge-Sphagnum community dominated by Carex
middendorffiiF.Schmidt, abundant, 4 September
2019, leg. N. Liksakova 17, det. E. Kuzmina (LE).
Sphagnum pulchrum is known from the oceanic
regions of the eastern part of North America, Europe,
Japan, and the Far East of Russia (Ignatov and Ignatova
2003). A fairly rare species for the southern Kuril
Islands, it was previously noted from the islands of
Iturup and Kunashir (Bakalin et al. 2009; Kuzmina and
Koroteeva 2016).
40. Sphagnum subfulvum Sjörs
Contributors. E. Yu. Kuzmina and N. S. Liksakova
Russia. Sakhalin Region, Urup Island, southern part of
the island, 6.02 km east of the mouth of the Kama
River, 45.6495168°N, 149.5463511°E, 244 m a.s.l., peat-
land with ridge-pool complexes, on the tussock, sedge-
Sphagnum community dominated by Carex midden-
dorffiiF.Schmidt with participation of dwarf-shrubs,
abundant and dominant, 4 September 2019, leg.
N. Liksakova 22, det. E. Kuzmina (LE).
This is a relatively rare species, known mainly from
the oceanic areas of the Holarctic Boreal Zone
(Ignatov and Ignatova 2003). Sphagnum subfulvum in
Russia has a mainly European distribution. Recently,
several localities for the species in the Asian part of
Russia have been found (Ellis, Alikhadzhiev, et al.
2020). In the South Kuril Islands it was previously
noted only from Shikotan Island (Bakalin et al. 2009).
41. Symphyogyna leptothelia Taylor
Contributor. E. Fuertes
Argentina. Prov. Jujuy, Dpto. Ledesma, Parque Nacio-
nal Calilegua, paraje Tres Cruces, epífita o en rocas
húmedas, en bosques montanos húmedos-hiperhú-
medos, las Yungas, de forófitos: Alnus acuminata
Kunth (aliso del cerro), Podocarpus parlatorei Pilg.
(pino del cerro), Cedrela angustifolia Sessé & Moc. ex
DC (cedro salteño), entre otros, 1060 m a.s.l., 23°44′S,
64°55′W, 16 June 2008, leg. E. Fuertes s.n. (MACB).
The plants are dioicous, olive-green, procumbent,
about 2–3cm long× 0.5–0.6 mm wide, dichotomously
forked, with a distinct midrib sharply defined from the
unistratose lamina, up to 1000 µm wide with one
strand of narrow, thick-walled cells. Thallus membranous,
undivided, margin with teeth 4–6 cells long, border
inconspicuous consisting of 1–2 rows of cells, about 62
(–95) long × 30(–35) µm wide, longer and narrower
than adjacent lamina cells, more or less isodiametric-hex-
agonal, about 70 (–75) long × 50 (–55) µm wide. Mucila-
ginous papillae present on the dorsal thallus surface.
Sporophytes scattered on the midrib, protected by tiny
scales. Seta long, whitish, capsule cylindrical, 4–5mm
long, dehiscing into 4 valves (Illustrations in Uribe and
Aguirre 1995,Figure7:A–D, Van Reenen 782).
204 L. T. ELLIS ET AL.
Symphyogyna leptothelia has been found as an epi-
phyte and on rocks, in moist and shaded deciduous
and evergreen forest at 800–1800 m a.s.l., in Jujuy pro-
vince (Argentina). Five species of Symphyogyna
are recognised in Argentina: S. aspera Steph. ex F.A.Mc-
Cormick, S. circinata Nees & Mont., S. podophylla
(Thunb.) Mont. & Nees, S. rubritincta A.Evans and
S. hochstetteri Nees & Mont. by Hässel de Menéndez
and Rubies (2009). Distributed in South America in
Colombia (Stephani 1900; Uribe and Aguirre 1995; Tro-
picos 2020), Perú (Stephani 1900; Uribe and Aguirre
1995), Brazil (Stephani 1900; Uribe and Aguirre 1995;
Gradstein and Costa 2003; Tropicos 2020) and Argen-
tina (this contribution).
42. Syntrichia latifolia (Bruch ex Hartm.) Huebener
Contributors. M. Boiko, N. Zagorodniuk and
L. M. Boiko
The Ukraine. Steppe zone, Kherson region, town Hola
Prystan’, park-monument of garden and park art of
local significance ‘Park of the “Hopry”Health Resort’,
on Ulmus laevis Pall., 1784, in the park near the
‘Soliane’lake, on bark at the base of trunks, associated
with Pylaisia polyantha (Hedw.) Schimp. (dominant epi-
phyte), Hypnum cupressiforme Hedw. and Leskea poly-
carpa Hedw., and at the soil surface Orthotrichum
diaphanum Schrad. ex Brid., 27 August 2020, leg.&
det. N. V. Zagorodniuk, M. Boiko s.n. (KHER).
Syntrichia latifolia is a xeromesophyte, eutrophic,
dioecious, nemoral species with a cushion-like turf
habit. The turfs are low (stems 2.5 cm high), blackish-
green and disintegrate easily; the leaves form apical
rosettes. In the sample studied capsules were not
found, but characteristic spherical, 2- and 4-celled
gemmae (mostly 2-celled) covered the leaf blades in
large quantities.
This species is known in the Ukraine from the far
west Zakarpattia region, Mukachevo district, the
Dobran’forestry (Zerov and Partyka 1975) and from
the far east Livoberezhnyi Lisostep, Kharkiv region,
Zmiivskyi district and the ‘Seredniodonetskyi’forest
reserve (Barsukov 2014). The present record was
found in the far south of the Ukraine and is a new
record for the steppe zone (Boiko 1992). Its locality is
significantly disjunct from most of the known range
of the species in southeastern Europe, which may be
explained by the mass formation of gemmae and
their ease of transfer, possibly by air masses, birds, or
humans. It is likely that dispersal to this location
occurred not long ago because the park was only
planted in the 1950s.
43. Syntrichia subpapillosissima (Bizot & R.B.Pierrot ex
W.Kramer) M.T.Gallego & J.Guerra
Contributor. H. Kürschner
Saudi Arabia. Asir Mountains, SW of Kamish Mushayt,
Jebel Natfa west of Al Harajah towards Al Jawwah,
17°55′21.79′′N, 43°16′32.24′′ E, 2800 m a.s.l., on rock in
Juniperus procera-Acacia origena woodland, 25 March
1984, leg. W. Frey & H. Kürschner 84-174,conf.
W. Kramer 2019 (herb. B, herb. Kürschner, Berlin).
Recorded as Tortula ruralis (Hedw.) P.Gaertn., E.Mey.
& Schreb. by Frey and Kürschner (1988). Syntrichia sub-
papillosissima belongs the heterogeneous S.ruralis
(Hedw.) F.Weber & D.Mohr complex (Kramer 1980).
Its morphological boundaries and taxonomic cat-
egories have been treated in different ways, but it
was elevated to the rank of a species by Gallego
et al. (2002). The species can be separated from
S. ruraliformis (Besch.) Cardot [S. ruralis var. ruraliformis
(Besch.) Delogn.] by the dense papillosity of the lamina
cells which is a ‘constant and sufficiently stable charac-
ter to support its status as a species’(Gallego et al.
2002).
A recent molecular and morphological analysis of
the Scandinavian Syntrichia ruralis complex revealed
that specimens with S. subpapillosissima morphology
are nested within S. ruralis or S. ruraliformis (Hedenäs
et al. 2019), the authors concluding that
S. subpapillosissima may represent only a specificphe-
notype, adapted to more arid conditions, of either
one of these two species. However, further studies
are required to investigate the taxonomic status of
the species (Hedenäs et al. 2019). A specimen from
the Asir Mountains of Saudi Arabia, previously
named Syntrichia ruralis (Frey and Kürschner 1988),
shows all of the morphological characters of
S. subpapillosissima [leaves densely papillose and
tapering towards apex, leaf apex (transition
between hairpoint and apex) hyaline, laminal cells
with pedicellate and branched papillae (bifurcate),
dorsal surface of costa strongly papillose to subden-
tate]. It is transferred to S. subpapillosissima and rep-
resents a new record for Saudi Arabia, increasing
the total number of bryophyte species for the
country to 129 (Taha 2019; Taha and Abou-Salama
in Ellis, Afonina, et al. 2020;KürschnerandFrey
2020a,2020b;Tahaetal.2020).
44. Tortella fasciculata (Culm.) Culm.
Contributors. P. Martin and S. V. O’Leary
Andorra. Serra Dels Corrals, on limestone outcrops
above communication mast, 42°33′20.6′′N, 1°27′02.9′′E,
2284 m a.s.l., 15 May 2019, close associates include
Tortella tortuosa (Hedw.) Limpr., Encalypta vulgaris
Hedw., Grimmia anodon Bruch & Schimp, Pseudocrossi-
dium obtusulum (Lindb.) H.A.Crum & L.E.Anderson,
Didymodon acutus (Brid.) K.Saito and Buckia vaucheri
(Lesq.) D.Rios, M.T.Gallego & J.Guerra. leg. & det.
P. Martin & S. V. O’Leary s.n., conf. T. L. Blockeel.
This is the first record for Andorra. Tortella fascicu-
lata is a suboceanic-submediterranean species and its
distribution in France and the Iberian Peninsula is
not completely clear. The confusion is due to the
JOURNAL OF BRYOLOGY 205
transfer of T.bambergeri (Schimp.) Broth. to either
T. fasciculata or T. pseudofragilis (Ther.) Köckinger &
Hedenäs (Köckinger and Hedenäs 2017).
In France it is known from the Pyrenees, from where
the type specimen originates, as well as in a few
localities in the Massif Central, Alsace and south-
eastern France (pers. comm. V. Hugonnot). The distri-
bution in Spain is unclear due to incomplete revision
of material, though there is a confirmed record from
Cuenca, Valdemoro Sierra, Cezon &Munoz (Köckinger
and Hedenäs 2017). Tortella fasciculata is not known
from Portugal (pers. comm. R. Porley).
45. Tortula cernua (Huebener) Lindb.
Contributors. S. Ursavaşand A. Özen
Turkey. Denizli, Honaz district, Aydınlar village, Kavaklı
creek, 37°44′24.90′′N, 29°22′33.49′′ E, 951 m. a.s.l., near
the creek bank (agricultural areas around the study
area),associated with Anomodon attenuatus (Hedw.)
Huebener, on soil, 15 May 2018, leg. A. Özen s.n., det.
S. Ursavaş, (U3277), conf. R.H. Zander.
Tortula cernua is reported here for the first time
from Turkey. It was recorded at the end of the 1900s
in South America (Cano and Gallego 2008), while
Bartram (1929) reported it in Canada from Sandy
Cove (Ingornachoix Bay), St. John Island, Bard Harbor
(St. John Bay) and Flower Cove (Tuomikoski et al.
1973). Although widely distributed on the North Amer-
ican continent, it is considered to be rare and endan-
gered in many European countries (Blanár and
Petrášová 2007), with conservation statuses such as
Regionally Extinct (RE) and Critically Endangered (CR)
(Hodgetts 2015; Hodgetts and Lockhart 2020). It is a
rare species over all its geographical range. It occurs
in unstable, calcareous, frequently transient natural
habitats, including on shores, soil, or rock, and in
anthropogenic habitats such as limestone quarries,
mines and walls (Hedenäs 2001). It is a boreal,
montane species, occurring mainly on calcareous soil,
frequently in maritime habitats. This record from
Turkey is the most southerly occurrence of T. cernua
in Europe and Asia. The presence of streams and lake
ecosystems in the region where it is located has
created a microclimatic effect (Ursavaşand Keçeli
2019) and allowed the species to spread in the area.
46. Tortula muralis Hedw.
Contributors. W. R. Álvaro Alba, D. A. Becerra Infante,
K. A. Cárdenas Espinosa and B. Moncada
Colombia. Boyacá, municipio de Tunja, Universidad
Pedagógica y Tecnológica de Colombia, 05°33′13.8′′ N,
73°21′41.3′′ W, 2730 m a.s.l., 19 January 2015, leg.
W. R. Álvaro Alba, D. A. Becerra Infante & K. A. Cárdenas
Espinosa 3425 (C-0019935 UDBC).
Tortula muralis (Pottiaceae) is a widely distributed
species (Zander 1993). The populations studied con-
sisted of small, yellowish-green plants with simple to
scarcely branching stems up to 0.4 cm long; leaves
crisped when dry, spreading when moist; lingulate;
apex rounded to obtuse; margins recurved from near
base to the apex, costa excurrent in hyaline hair-
point; medial and upper cells pluripapillose with bifur-
cated papillae; seta erect, twisted, yellow to yellowish
brown; calyptra cucullate; capsule erect, cylindrical;
operculum conical. This species was found in an
urban habitat, on brick and concrete walls, in
exposed and humid places, growing in short turfs. In
the most recent treatment of the genus for South
America, Cano and Gallego (2008) state that the
species should be excluded from the Neotropics,
after checking specimens collected in Peru actually
correspond to Syntrichia limensis (R.S.Williams)
R.H.Zander. However, taking into account that the
species tends to establish itself on artificial substrates,
it can be inferred that the scarce collections of urban
bryophytes in the Neotropical countries do not allow
us to surmise the real distribution of the species.
47. Ulota intermedia Schimp.
Contributor. A. Graulich
Belgium. (1) Marchin, Hoyoux valley, on Fagus sylva-
tica L., 50°29′20′′N, 5°15′17′′ E, 210 m a.s.l., 18 January
2020, leg. A. Graulich, det A. Graulich (herb. Graulich
ULOTA 09/20); (2) Malmedy, Moûpa, on Fagus sylvatica,
50°28′08′′N, 6°01′50′′ E, 550 m a.s.l., 28 June 2020, leg.
A. Graulich, det. A. Graulich (herb. Graulich ULOTA
18/20).
In Europe, Ulota intermedia has its main distribution
area in the northern and central part of the continent.
In southern Europe, the species is restricted to
montane areas (Caparrós et al. 2016). Nevertheless,
its distribution is still imperfectly known due to its
new taxonomic status, adding to the confusion with
other epiphytic species such as U. bruchii Hornsch.
and U. crispula Bruch. Ulota intermedia is probably
widespread in the hilly area of eastern Belgium.
48. Ulota pycnophylla Dusén ex Malta
Contributors. R. Garilleti, B. Albertos, I. Draper and
F. Lara
Argentina. Neuquén, Departamento Los Lagos,
Parque Nacional Nahuel Huapi, Lake Pireco, Cardenal
Samoré International pass, near the Argentinian
customs post, 40°43’16′′S, 71°47’57′′ W, 800 m a.s.l.,
Coihue tree (Nothofagus dombeyi Bl.) forest,epiphyte
on young N. dombeyi, 13 January 2017, leg.
R. Garilleti 2017-095a, 096e with F. Lara.
This moss was described (Malta 1927) based on a
specimen collected by Dusén from Cordillera de la
Costa, near Angol (Araucanía Region), an area with a
Mediterranean climate. It was not found again until
its rediscovery far south at Capitan Prat Province,
Aysén Region (Larraín 2016), a zone with a temperate
oceanic climate. This is the first record for Argentina
206 L. T. ELLIS ET AL.
and the easternmost one, representing an intermedi-
ate point between the extremes of the known distri-
bution of the species, while also being from an area
with an oceanic climate.
Larraín (2016) pointed out that Ulota pycnophylla
could be a synonym of U. macrocalycina Mitt., but
with some differences that could sustain its distinction.
Both species share characters such as a creeping
growth form and leaves scarcely curved to straight
with a (sub-)obtuse apex. Discriminating characters
are: U. macrocalycina has broader leaf apices, short
and dense branches, naked calyptrae and spores 30–
35 μm in diameter; U. pycnophylla has obtuse to
acute leaf apices, longer lax branches, hairy calyptrae
and spores 20–26 μm. Ulota macrocalycina shows a
preference for hyper-humid oceanic cold climates. It
is widespread in Tierra del Fuego (Matteri 2003;
Larraín and Bahamonde 2017), and has rarely been
recorded from more northerly areas such as Aysén
(Larraín 2016) and Los Lagos regions (Deguchi 1991).
The latter record is based on a specimen from the
Chilean–Argentinian border, very close to our own
finding of U. pycnophylla in Argentina. It has not
been possible for us to study that specimen to
confirm the identification.
Acknowledgements
The work of V. A. Bakalin, S. S. Choi and C. W. Hyun was sup-
ported by the grant ‘Survey of Korean Indigenous Species’
from the National Institute of Biological Resources of Ministry
of Environment in Korea awarded to C. W. Hyun. Field work of
M. S. Ignatov and E. A. Ignatova was supported by RFBR
19-04-00976. The study by A. Uygur, T. Ezer, S. Karaman
Erkul and M. Alataşwas supported financially by the
Scientific and Technological Research Council of Turkey
(TÜBİTAK) (project number 120Z046), which is thanked for
its financial support.
E. Fuertes thanks the Argentine National Parks for provid-
ing her with collecting permits. Her research was funded by
project AECI, A- 8930-2007 of the Science and Innovation Min-
istry of Spain. H. Kürschner is grateful to Dr W. Kramer (Frei-
burg) for the revision of some Tortula/Syntrichia species of
the S. ruralis complex. María J. Cano thanks Jaime Güemes
and Leopoldo Medina for the organisation of the botanical
expedition to Crete and Manolis Avramakis and the staffof
the Natural History Museum of Crete (NHMC) for support
and logistics. This work was financed by the Spanish Ministerio
de Ciencia e Innovación (project PID2019-103993GB-I00 co-
financed by FEDER). Pamela Saha, Md. Nehal Aziz and Debab-
rata Maity thank the former Director of the Botanical Survey of
India, Dr. Paramjit Singh, and the present Director, Dr. A. A.
Mao, for providing necessary facilities and encouragement.
They are also grateful to PCCF, West Bengal for granting per-
mission for the field exploration tours. The first author (P.
Saha) thanks the Principal and Head of the Department of
Botany, Bijoy Krishna Girl’s College, Howrah, West Bengal, for
encouragement and necessary facilities.
K. K. Rawat, V. Sahu and R. R. Paul are grateful to the Direc-
tor of the CSIR-National Botanical Research Institute for
kindly providing the facilities required for the study. They
are also grateful to the forest department officials of
Tawang, Arunachal Pradesh and Tungnath, Uttarakhand for
kindly permitting the study. Financial assistance from the
Space Application Centre, ISRO, Ahmedabad to KKR and
RRP under the project number GAP-3464 is also acknowl-
edged. The authors also thank the institutional ethics com-
mittee for granting institutional MS number CSIR-NBRI_MS/
2020/12/02. The contribution of Michaela Kropik and
Harald Zechmeister is part of the LE 14–20 project ‘Natur-
raum Nationalpark Thayatal 2020+ (7.6.1b-II6-47/19)’. The
authors acknowledge the financial support of their study
by the Nationalpark Thayatal GmbH. The work of
E. A. Borovichev and N. R. Shafigullina was partially sup-
ported by the Russian Foundation of Basic Researches
(grant number 18-04-00594A). S. Ștefănuțacknowledges
the support of project number RO1567-IBB03/2020 through
the Institute of Biology Bucharest of the Romanian
Academy. The work of M. V. Dulin was performed within
the scope of the state task ‘Diversity of vegetation at the
west macroslope of the Subpolar Ural’(number АААА-
А19-119011790022-1). The author is also grateful to
A. D. Potemkin for confirming the identity of Scapania
kaurinii.
The study by A. D. Potemkin was carried out within the
framework of the institutional research project of Komarov
Botanical Institute of the Russian Academy of Sciences
‘Flora and systematics of lichens and bryophytes of Russia
and phytogeographically important regions’(АААА-А19-
119020690077-4) and was supported by RFBR project 18-
05-60093 Арктика. The fieldwork of M. Ryan was supported
by the Polar Continental Shelf Program (Natural Resources
Canada), the Yukon Department of Environment, and
Environment and Climate Change Canada. M. Ryan and
A. D. Potemkin are grateful to T. McIntosh for collecting Sca-
pania obscura.M.Kırmacı, F. Filiz and U. Çatak cordially thank
TUBİTAK for financial support to Mesut Kırmacı(TBAG
113Z631), and also thank Harald Kürschner for kind help
during field trips and with regard to the genus Sphagnum.
E. Fuertes expresses her gratitude to Olga Martínez and
Lázaro J. Novara from the University of Salta (Argentina),
for the attention and facilities received for the study and con-
servation of the bryological material. Her research was
funded by the Spanish Foreign Office of the Kingdom of
Spain, project AECI, A /8930/2007.
J. G. Segarra-Moragues thanks P. Ballings and A. Bogaerts
(Botanic Garden Meise Herbarium), for their help in locating
and accessing specimens, and F. Puche for comments on an
earlier version of their note. Their study was partially funded
by the Spanish Ministry of Economy, Industry and Competi-
tiveness project CGL2016-80418-P. S. Ursavaşand A. Özen
would like to thank Richard H. Zander for his help in confi-
rming the determination of Tortula cernua. The contribution
by R. Garilleti, B. Albertos, I. Draper and F. Lara was supported
by the Spanish Ministry of Economy and Competitiveness
grant CGL2013–43246-P and Economy, Industry and Compe-
titiveness grant CGL2016-80772-P. The authors acknowledge
the Argentinian Administración de Parques Nacionales for
authorisation to collect bryophytes in the territories
managed by this national service.
The work reported by V. E. Fedosov was supported by RSF
grant # 18-14-00121. J. Enroth thanks Dr. Matti Nummelin
(Hyvinkää, Finland) for sending the specimens for identifi-
cation. The work of N. A. Konstantinova and A. N. Savchenko
was conducted within the framework of the institutional
research project АААА-А18-118050490088-0 and was partly
supported by RFBR grants 18-04-00594 and 18-05-60093.
The study reported by E. Yu. Kuzmina was carried out
within the framework of the basic project (planned theme
JOURNAL OF BRYOLOGY 207
of the laboratory of lichenology and bryology of Komarov
Botanical Institute ‘Flora of lichens and bryophytes of
Russia and phytogeographically important regions’)
number АААА-А19-119020690077-4 and partially sup-
ported by the grant of the Russian Foundation for Basic
Research number 19-05-00805-а. The study of N. S. Liksakova
was carried out within the framework of the planned theme
of the Laboratory of Geobotany of the Komarov Botanical
Institute ‘Vegetation of European Russia and Northern Asia:
diversity, dynamics and organization principles’number
121032500047-1 N. S. Liksakova express her gratitude to
the organisers of and participants in the expedition to the
Kuril Islands in 2019 ‘East Bastion –Kuril Ridge’, organised
by the Expedition Center of the Ministry of Defense of the
Russian Federation and the Russian Geographical Society.
The contribution by G. F. Peñaloza-Bojacá and S. A. Maciel-
Silva was financed in part by Coordenação de Aper- feiçoa-
mento de Pessoal de Nível Superior-Brasil (CAPES)- Finance
Code 001, and the Rufford Foundation (small grant 2018-
2019). W. R. Álvaro Alba, D. A. Becerra Infante, K. A. Cárdenas
Espinosa and B. Moncada thank the Corporación Autónoma
Regional de Boyacá (Corpoboyacá) for granting the study
permit for scientific research in biological diversity with
permit number 3524 of 2014, and María Teresa Gallego for
the confirmation of Tortula muralis.S.Ștefănuțacknowledges
the support by project number RO1567-IBB03/2021 through
the Institute of Biology Bucharest of the Romanian Academy.
ORCID
V. A. Bakalin http://orcid.org/0000-0001-7897-4305
I. Draper http://orcid.org/0000-0003-3102-9386
R. Garilleti http://orcid.org/0000-0002-5977-2908
B. Albertos http://orcid.org/0000-0002-2116-5600
S. Sciandrello http://orcid.org/0000-0003-1132-5698
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