Arctoa (2017) 27: 49-53
doi: 10.15298/arctoa.27.05
ON THE GENUS LESKEA (LESKEACEAE, BRYOPHYTA) IN RUSSIA
РОД LESKEA (LESKEACEAE, BRYOPHYTA) IN RUSSIA
MICHAEL S. IGNATOV1,2, ANDREY G. BEZGODOV3, OXANA I. KUZNETSOVA2 & ELENA A. IGNATOVA1
МИХАИЛ С. ИГНАТОВ1,2, АНДРЕЙ Г. БЕЗГОДОВ3, ОКСАНА И. КУЗНЕЦОВА2, ЕЛЕНА А. ИГНАТОВА1
Abstract
The genus Leskea in Russia is represented by one widespread species, L. polycarpa Hedw. Records
of L. gracilescens appeared to be erroneous. Analysis of nuclear ITS sequence data demonstrated that
North American samples of L. gracilescens and L. obscura clearly differ from European and Asian
populations in several characteristic substitutions. Anomalous morphology of plants from Yakutia and
Amur Province is described and illustrated. Their distribution likely corresponds to a high and late
floodings of big rivers, crossing permafrost areas.
Резюме
Род Leskea представлен в России одним широко распространенным видом , L. polycarpa Hedw.
Указания на находки L. gracilescens в Амурской области были ошибочными. Анализ нуклеотидных
последовательностей ядерного спейсера ITS показал, что североамериканские образцы L. gracilescens and L. obscura отличаются несколькими заменами от европейских и азиатских растений.
Необычная морфология растений L. polycarpa из Амурской области и Якутии, вероятно, связана
с сильными поздними паводками на больших реках, протекающих в зоне вечной мерзлоты.
KEYWORDS: Leskea, Russian Far East, Yakutia, ecomorphs
INTRODUCTION
The genus Leskea in Russia included just one species,
L. polycarpa Hedw., in almost all publications (Abramov
& Volkova, 1998; Abramova et al., 1961; Ignatov,
Afonina, Ignatova et al., 2006). Although Hedwig (1801)
described from Europe two species, L. polycarpa and L.
paludosa, the latter one was considered as a variety of
the former already by authors of 19th century (e.g.
Schimper, 1860). Eleven infraspecific taxa within Leskea
polycarpa are listed in Torpicos database (http://
www.tropicos.org/namesearch.aspx, accessed 1 May
2018), but in recent checklists (Hill et al., 2006; Ros et
al., 2013) and Floras (Brugués & Ruiz, 2018; Redfearn,
2014) they are usually not mentioned at all. Leskea
latifolia Lindb. ex Broth., described from the Caucasus
(Brotherus, 1892), was synonymized with Amblystegium
varium (Abramov, 1969).
In the course of the flora inventory studies in the Nature Reserve «Norsky», Amur River basin in the Russian
Far East, we recently reported Leskea gracilescens (Bezgodov et al., 2013), as these plants were sufficiently different, agreeing in many characters with this species as
it was circumscribed in moss floras of North America
(Redfearn, 2014; Crum & Anderson, 1981) and espe1
cially moss flora of China (Cao et al., 2002). In China
this species was reported only recently from its NE part,
on the right bank of Amur River in a relatively close
proximity to the area where the mentioned Russian collection has been done. Chinese collections were reported
as sterile (Cao et al., 2002), while in Russian specimens
sporophytes were immature, thus the sporophyte characters were impossible to use. However, catenulate foliage,
smaller stem leaves with the smaller length to width ratio, straight and symmetric leaves, lacking characteristic
obligue apical part, and occasionally blunt apices in both
stem and branch leaves fitted the description of L. gracilescens.
Later in Yakutia, to the north of Amur Province, we
collected Leskea with even more obtuse leaves, more similar to another American species, L. obscura. So we decided to test with DNA markers if the plant from Amur
Province and Yakutia really belong to L. gracillescens or
represent another taxon.
MATERIALS AND METHODS
Sampling. In addition to L. gracillescens-like plants,
we included the most common phenotypes of L. polycarpa from the Russian Far East and Yakutia into the set
for sequencing, and specimen from European Russia.
– Lomonosov Moscow State University, Biological Faculty, Geobotany Dept., Leninskie Gory Str. 1-12, Moscow 119234 Russia
– Россия, 119234, Москва, Ленинские Горы, д. 1 стр. 12, Московский государственный университет, биологический
факультет, кафедра геоботаники. E-mails: misha_ignatov@list.ru & arctoa@list.ru
2 – Tsitsin Main Botanical Garden, Russian Academy of Sciences, Botanicheskaya Str., 4, Moscow 127276 Russia – Россия
127276 Москва, Ботаническая 4, ГБС РАН. E-mail: oikuznets@gmail.com
3 – Joint Stock Company Kama Research Institute on Complex Deep and Superdeep Borehole Investigations, Krasnoflotskaya str.,
15, Perm, 614016 Russia. – Россия 614016, Пермь, Краснофлотская, 15, АО «Камский научно-исследовательский институт
комплексных исследований глубоких и сверхглубоких скважин». E-mail: bezgodovs@list.ru
50
M.S. IGNATOV, A.G. BEZGODOV, O.I. KUZNETSOVA & E.A. IGNATOVA
100 µm
5
2
3
1 cm
2 mm
4
6
9
7
8
1
10
13
1 mm
15
14
11
12
16
17
18
Fig. 1. Leskea polycarpa (from: Russia, Yakutia, Khangalassky Distr., 2.VIII.2016, Ignatov & Ignatova #16-160, MHA). 1, 4
– habit, dry; 2–3 – capsules; 5 – leaf transverse section; 6 – stem transverse section; 7–9 – paraphyllia; 10 – upper laminal cells;
11 – basal laminal cells; 12 – median laminal cells; 13–15 – stem leaves; 16–18 – branch leaves. Scale bars: 1 cm for 4; 2 mm for
1–3; 1 mm for 13–18; 100 µm for 5–12.
On the genus Leskea in Russia
51
100 µm
10
9
2
3
12
8
1
4
11
1 mm
14
6
5
15
13
7
17
18
19
22
16
20
21
24
23
25
32
29
27
26
28
30
31
33
Fig. 2. 1–5 – Leskea polycarpa (1–7 from: Russia, Amursakaya Prov., 1.VII.2010, Bezgodov #34, MHA; 8–15 – from: Russia,
Sakhalin, Pisarenko #0p03765, MHA); 16–21 – L. gracilescens (from: U.S.A., Ohio, Buck #50264, NY); 22–25 – L. obscura
(from: U.S.A., Buck 22206, NY); 26–33 – L. cf. gracilescens (from: Russia, Amurskaya Prov., 1.VII.2010, Bezgodov #210, MHA).1,
6, 16, 22, 26, 28– upper laminal cells; 2–3, 9–10, 20–21, 32–33 – branch leaves; 5–7, 13–15, 17–19, 23–25, 29–31 – stem leaves;
4, 11–12, 27 – paraphyllia. Scale bars: 1 mm for 2–3, 5–7, 9–10, 12–15, 17–21, 23–25, 29–33; 100 µm for 1, 4, 8, 11–12, 16, 22,
26–28.
52
M.S. IGNATOV, A.G. BEZGODOV, O.I. KUZNETSOVA & E.A. IGNATOVA
Fig. 3. Five fragments of the ITS alignment of Leskea, showing characteristic substitutions indicating identity of Russian plants
of different morphotypes in costrast to American specimens of L. gracilescens and L. obscura. Specimens details are in Table 1
and complete aliggnment in Supplementary materials.
Molecular protocols.
DNA extraction and amplification were done according to the laboratory protocols described in Gardiner et
al. (2005). Vouchers specimens and GenBank accession
numbers are listed in Table 1. Sequences were aligned
manually in BioEdit (Hall, 1999).
RESULTS
The molecular results are partly shown in Fig. 3, and
complete alignment is given as a supplementary material. There are only very few substitutions between American samples of L. gracilescens and L. obtusa and Eurasian plants, thus we present their distinction by a parts
of alignment with substitutions, instead of a tree or haplotype network. The ITS region is found to be variable
for separation of these three entities by characteristic substitutions. Among them, L. gracilescens and L. polycarpa
lack infraspecific variation in these positions, and L.
obtusa has one vatiable position.
Specimens from the Amur Province, as well as from
Yakutia revealed no differences in sequences from specimens of the common phenotypes of L. polycarpa, both
from the Far Eastern and European populations (Fig. 3).
DISCUSSION
Although the present study results in no taxonomic
novelties, the fact of such extreme variation is worthy
describing and illustrating (Figs. 1–2). Obviously, the
record of L. gracilescens from Russia and probably also
from China are based on the ecomorphs of L. polycarpa.
Leaf apices in L. polycarpa are described as ‘blunt or
acute’ in “Moss Flora of North America” (Redfearn Jr.,
2008); ‘acute to obtuse’ in “The moss flora of Britain
and Ireland” (Smith, 2004); ‘agudo o acuminado’ in
“Flora bryofítica Ibérica” (Brugués, 2018); “blunt to
acute” in “Moss flora of the Middle European Russia”
(Ignatov & Ignatova, 2004). Illustrations in all these floras show slightly secund leaves with more or less oblique apices, only rarely straight and symmetric. This
character, along with differentiated stem and branch
leaves (stem leaves being longer than wide), is considered as important for separating of L. polycarpa from L.
gracilescens by Redfearn Jr. (2008): L. gracilescens, contrary to L. polycarpa, has straight leaves with symmetrical apices, as well as stem and branch leaves equally
hardly longer than wide.
Most Leskea specimens from Russia fully agree with
the mentioned features of L. polycarpa, except for the
anomalous specimen from Amurskaya Province referred
previously to L. gracilescens. In Fig. 2: 1–15, two specimens of L. polycarpa from Asian Russia are shown, with
clearly asymmetrical stem leaves, oblique leaf apices and
less oblong branch leaves with obtuse apices. However,
specimen from Amurskaya Province (Fig. 2: 26–33) resembles L. gracilescens from North America (Fig. 2: 16–
21) rather than L. polycarpa in shape of stem and branch
leaves, which are straight and have symmetrical apices.
Nevertheless, evidence from molecular markers suggests
that this deceiving resemblance is likely caused by the
influence of ecological conditions.
Another peculiar specimen with all leaves obtuse, even
Table 1. Specimens of the genus Leskea used in molecular study, with GenBank accession numbers.
Species
L. gracilescens
L. gracilescens
L. gracilescens
L. obtusa
L. obtusa
L. obtusa
L. polycarpa
L. polycarpa
L. polycarpa
L. polycarpa
L. polycarpa
L. polycarpa
Specimen
USA: Connecticut, coll. Tan #91502 (MHA)
USA: Chicago, 6.X.2013 Ignatov (MHA)
USA: New York, Buck 57814 (NY barcode 1221900)
USA: New Jersey, Buck 22206 (NY barcode 489277)
USA: Missouri, Buck 30103 (NY barcode 489398)
USA: Kansas, Buck 46377(NY barcode 680473)
Russia: Amurskaya Province, 1.VII.2010 #69 (MHA)
Russia: Ryazan, 11.XI.2009 Donskov (MHA)
Russia: Amurskaya Province, 1.VII.2010 Bezgodov #34 (MHA)
Russia: Vladivostok, Ignatov #08-355a (MHA)
Russia: Sakhalin, Pisarenko #3765 (MHA)
Russia: Yakutia, Ignatov & Ignatova 16-160 (MHA)
Isolate
OK555
OK556
OK541
OK545
OK544
OK542
OK428
OK551
OK552
OK553
OK554
OK1438
Genbank number
MH558262
MH558263
MH558264
MH558265
MH558266
MH558267
MH558268
MH558269
MH558270
MH558271
MH558272
MH558273
On the genus Leskea in Russia
with rounded apices, was collected in Central Yakutia.
However, its stem leaves are more longer than wide in
comparison with branch leaves and have slightly oblique
apices (Fig. 2), which is in agreement with circumscription of L. polycarpa according to Redfearn Jr. (2008).
Capsules in this specimen are also slightly different from
common morphotype of L. polycarpa, being straight and
shorter than usual. However, molecular markers did not
reveal any difference between this specimen and other
specimens of L. polycarpa, pointing on wider morphological variability of this species in peculiar ecological
conditions in permafrost areas of Asian Russia with repeated strong summer floods caused by the permafrost
melting.
ACKNOWLEDGEMENTS
The work was supported by RSF (grant no. 18-1400121).
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