New Taxonomic Arrangement of Dicranella s.l. and Aongstroemia s.l. (Dicranidae, Bryophyta)

Abstract

The recent molecular phylogenetic study of the families Aongstroemiaceae and Di-cranellaceae, which resolved the genera Aongstroemia and Dicranella as polyphyletic, indicated the need for changes in their circumscription and provided new morphological evidence to support the formal description of newly recognized lineages. Following up on these results, the present study adds another molecular marker, the highly informative trnK-psbA region, to a subset of previously analyzed taxa and presents molecular data from newly analyzed austral representatives of Di-cranella and collections of Dicranella-like plants from North Asia. The molecular data are linked with morphological traits, particularly the leaf shape, tuber morphology, and capsule and peristome characters. Based on this multi-proxy evidence, we propose three new families (Dicranellopsida-ceae, Rhizogemmaceae, and Ruficaulaceae) and six new genera (Bryopalisotia, Calcidicranella, Di-cranellopsis, Protoaongstroemia, Rhizogemma, and Ruficaulis) to accommodate the described species according to the revealed phylogenetic affinities. Additionally, we amend the circumscriptions of the families Aongstroemiaceae and Dicranellaceae, as well as the genera Aongstroemia and Di-cranella. In addition to the monotypic Protoaongstroemia that contains the newly described di-cranelloid plant with a 2-3-layered distal leaf portion from Pacific Russia, P. sachalinensis, Dicranella thermalis is described for a D. heteromalla-like plant from the same region. Fourteen new combinations , including one new status change, are proposed.

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Plants 2023, 12, 1360. https://doi.org/10.3390/plants12061360 www.mdpi.com/journal/plants
Article
New Taxonomic Arrangement of Dicranella s.l. and
Aongstroemia s.l. (Dicranidae, Bryophyta)
Vladimir Fedosov 1,2,*, Alina Fedorova 3, Elena Ignatova 1 and Jan Kučera 4
1 Biological Faculty, Lomonosov Moscow State University, Moscow 119234, Russia
2 Botanical Garden-Institute, FEB RAS, Vladivostok 690024, Russia
3 Main Botanical Garden, Russian Academy of Sciences, Moscow 127276, Russia
4 Department of Botany, Faculty of Science, University of South Bohemia,
CZ–370 05 České Budějovice, Czech Republic
* Correspondence: fedosov_v@mail.ru
Abstract: The recent molecular phylogenetic study of the families Aongstroemiaceae and Di-
cranellaceae, which resolved the genera Aongstroemia and Dicranella as polyphyletic, indicated the
need for changes in their circumscription and provided new morphological evidence to support the
formal description of newly recognized lineages. Following up on these results, the present study
adds another molecular marker, the highly informative trnK–psbA region, to a subset of previously
analyzed taxa and presents molecular data from newly analyzed austral representatives of Di-
cranella and collections of Dicranella-like plants from North Asia. The molecular data are linked with
morphological traits, particularly the leaf shape, tuber morphology, and capsule and peristome
characters. Based on this multi-proxy evidence, we propose three new families (Dicranellopsida-
ceae, Rhizogemmaceae, and Ruficaulaceae) and six new genera (Bryopalisotia, Calcidicranella, Di-
cranellopsis, Protoaongstroemia, Rhizogemma, and Ruficaulis) to accommodate the described species
according to the revealed phylogenetic affinities. Additionally, we amend the circumscriptions of
the families Aongstroemiaceae and Dicranellaceae, as well as the genera Aongstroemia and Di-
cranella. In addition to the monotypic Protoaongstroemia that contains the newly described di-
cranelloid plant with a 2–3-layered distal leaf portion from Pacific Russia, P. sachalinensis, Dicranella
thermalis is described for a D. heteromalla-like plant from the same region. Fourteen new combina-
tions, including one new status change, are proposed.
Keywords: Haplolepidous mosses; nad5 G1 intron; phylogenetics; polyphyly; morphological
convergence; trnS-trnF region; trnK; biodiversity
1. Introduction
Molecular phylogenetic studies of bryophytes [1–10] have shown that the traditional
morphology-based circumscriptions of genera are often biased by morphological conver-
gence and in reality comprise a suite of phylogenetically unrelated lineages. This is espe-
cially the case for genera with reduced morphology, such as pioneer species with short
life cycles, e.g., Entosthodon Schwägr. ex Hornsch., Physcomitrium (Brid.) Brid. [11,12], or
Ditrichum Timm ex Hampe [8], although the larger pleurocarpous mosses, such as in the
traditionally delimited Hygrohypnum or Hypnum, have also been shown to be prone to the
homoplasic retention of distinct morphological features in unrelated lineages [7,13–15].
It was thus perhaps not very surprising that a molecular-phylogenetic study of the
northern temperate genera of Aongstroemia Bruch & Schimp. and Dicranella (Müll. Hal.)
Schimp. [10] revealed a striking polyphyly in the existing delimitation of these genera.
The traditional morphological circumscription of the genus Dicranella included plants of
small size with a stem central strand, elongated linear-lanceolate to subulate leaves, costae
Citation: Fedosov, V.; Fedorova, A.;
Ignatova, E.; Kučera, J. New
Taxonomic Arrangement of
Dicranella s.l. and Aongstroemia s.l.
(Dicranidae, Bryophyta).
Plants 2023, 12, 1360.
https://doi.org/10.3390/plants12061360
Academic Editor: Yin-Long Qiu
Received: 29 January 2023
Revised: 6 March 2023
Accepted: 14 March 2023
Published: 17 March 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license
(https://creativecommons.org/license
s/by/4.0/).

Plants 2023, 12, 1360 2 of 40
with guide cells in the cross-section, predominantly dioicous sexual condition, and dicra-
noid peristome [16–20]. This allowed for a considerable variability of the other gameto-
phytic and sporophytic traits, which found a reflection in the molecular differentiation of
the thirteen analyzed species of the genus that were found in seven different phylogenetic
lineages of haplolepidous mosses (the subclass of Dicranidae), three of which could be
considered orphaned in the system of the representatives of dicranids analyzed to date.
On the other hand, the genus Aongstroemia, originally introduced for a single species, A.
longipes, was soon substantially expanded [16] to include most species of the modern Di-
cranella s.lat., and later again reduced [21] to harbor species sharing the julaceous habit
originating from the ovate leaves, which are broadly rounded to acute or slightly attenu-
ate at the tips and appressed to the stem. Despite this restriction in the generic concept of
Aongstroemia, the rate of cryptic molecular diversity was similar to that revealed in Di-
cranella; Bonfim-Santos et al. [10] showed that the three analyzed species (out of the 11
accepted names in the genus) appear in three lineages belonging to three currently recog-
nized families. Although the polyphyly of both genera was demonstrated quite convinc-
ingly, this study has not yet resulted in a taxonomic treatment. The major reason for this
was the insufficient taxonomic sampling, particularly in Dicranella: of the 161 accepted
species according to the Tropicos database [22], plus the 47 accepted species in Leptotrich-
ella and five accepted names in Anisothecium, less than one tenth have been phylogenet-
ically studied, which means that the generic and familial assignment of the bulk of the
species remains pending after the splitting of Dicranella according to the obtained results.
The recent description of a previously unknown dicranelloid moss from SW Portugal,
which necessitated the erection of a new genus, Neodicranella Porley & Fedosov, following
the assessment of molecular affinities [23], confirms that the diversity of dicranelloid
mosses has not been fully captured, even in the relatively well-surveyed Europe. Alt-
hough a thorough taxonomic revision of all included taxa and checking of the type mate-
rial would be most appropriate, such a revision will hardly be possible in the near future
for a complex and species-rich genus such as Dicranella, where many of the accepted spe-
cies are based on a few historical collections from southern tropical countries, whose lo-
calities are difficult to access. Nomenclaturally, it would nevertheless be more relevant if
the types of the generic names placed earlier in synonymy with Dicranella and Aongstroe-
mia were designated, and their phylogenetic affinities known; a task that has been partly
accomplished and the missing pieces of information do not threaten the stability of most
proposed taxonomic solutions. Moreover, further accepting clearly polyphyletic taxa is in
our opinion a less desirable alternative than establishing a baseline for further develop-
ment of a phylogenetically-based system of haplolepidous mosses, which can be further
elaborated as soon as new information appears.
Although we generally followed the molecular sampling of Bonfim Santos et al. [10],
who employed only organellar markers, plastid trnL–trnF and rps4 and mitochondrial
nad5 intron 2, with respect to the absence of reasonably informative nuclear markers that
would be generally used in subclass-spanning phylogenetic studies of mosses, we deep-
ened the molecular sampling to include the highly informative trnK–psbA region, which
has been used with success, e.g., in the treatments [24] or [25], and also sampled the two
variable spacers flanking the gene for tRNA-Thr, which is located between the previously
sampled trnL–trnF and rps4. The purpose of this was the testing of the weakly supported
deeper nodes of Dicranidae, which was one of the unresolved questions in [10] that we
aimed to address in our novel analyses. Secondly, we broadened the sampling in several
critical groups, such as the South American representatives of Dicranella assigned to Ani-
sothecium by Mitten [26], accessions of Neodicranella and several putatively new taxa of
unclear affinity from North Asia, which were not included in [10]. We also deliver argu-
ments for treating the two varieties of Dicranella schreberiana (Hedw.) Hilf. ex H.A. Crum
& L.E. Anderson at the species level and resolve the molecular variation between D. varia
(Hedw.) Schimp. and D. howei Renauld & Cardot. This study thus represents a state-of-

Plants 2023, 12, 1360 3 of 40
the-art taxonomic treatment based on both previously published and newly obtained mo-
lecular and morphological data, which is expected to be updated, particularly for the
southern and tropical taxa.
2. Results
The concatenated matrix consisted of 6381 aligned sites, of which 563 belonged to L
partition, 975 to T partition, 728 to R, 2899 to K, and 1216 to N. Indels scored for the L, R,
and N partitions yielded an additional 193 binary sites, and 251 indels were scored for the
T and 218 for the K partition. The partitions corresponding to the dataset used by Bonfim-
Santos et al. [10], i.e., L, R, and N, contained 907 variable and 603 parsimony-informative
sites, the T partition had 526 variable and 346 parsimony-informative sites, and the K par-
tition contained 1410 variable and 901 parsimony-informative sites.
The trees inferred from the combined L, R, and N data, and those with an added T
region dataset, had essentially identical topology, with generally higher support values
from the expanded dataset. The addition of indel data generally further improved the
support values, without changes in topology at the supported nodes, however only when
T indel data were not considered. At the same time, the trees estimated from the separate
analysis of K data, which again had generally higher support node values in the version
with SIC-coded indels included, yielded a topological incongruence compared to the trees
derived from LTRN data, with respect to the estimated affinities of Chrysoblastella chilensis,
Neodicranella hamulosa, and Archidium + Leucobryaceae and Grimmiales clades. Hence, we
present here the results representing the total evidence of the fully concatenated dataset
(LTRKN) in Figure 1, and the trees resulting from partial analyses of the LTRN and K data
are presented as the supplementary Figures S1 and S2. In the following description of the
results, we only comment on results differing from those obtained by Bonfim-Santos et al.
[10], which was used as a reference.

Plants 2023, 12, 1360 4 of 40
Figure 1. Maximum likelihood tree, inferred from the concatenated data matrix from the chloroplast
trnF–trnS and trnK–psbA and mitochondrial nad5 intron 2 sequence data (LTRKN dataset) of se-

Plants 2023, 12, 1360 5 of 40
lected species of Dicranidae, focused on the genera Dicranella and Aongstroemia, rooted with Pseudo-
ditrichum mirabile. Bootstrap support values higher than 60 inferred from ML analyses, without and
with indel coding, and posterior probabilities higher than 0.7 inferred from BI, without and with
indel coding, are shown above the branches; hyphens in place of support values denote lower sup-
port of the node, while a blank space indicates that the node is absent from the topology inferred
from the particular analysis; maximally supported nodes are indicated by solid lines and asterisks.
Newly studied terminals, as well as terminals for which at least one marker was obtained de novo,
are printed in bold, and terminals for which the trnK–psbA sequence is available are underlined. For
details, see Appendix A.
In contrast to this study, after reconsidering the reading of several bases in the raw
chromatograms, the position of Pseudoditrichum mirabile was newly assessed as unre-
solved among the basal protohaplolepidous clades (Flexitrichum, Scouleria + Drummondia
+ Hymenoloma clade) and the clade containing all other analyzed taxa, i.e., Bryoxiphium +
rest of Dicranales incl. Grimmiales), rather than being found in the basal grade of Di-
cranales. This position was shared by the analyses resulting from the LTRN and K datasets
(Figures S1 and 2). Additional Dicranella staphylina accessions from northeastern Asia
(Putorana, Yakutia, and Khabarovsk Territory) were found in the maximally supported
clade with European accessions, although three accessions collected in the heart of the
permafrost zone of northeast Asia (BF59, FDt107, and 116) differed in several substitu-
tions, despite the absence of notable morphological differences, except for the slightly
more robust habit. European accessions of the previously unanalyzed D. humilis from the
Czech Republic and Russian Leningrad Province proved identical and distinct from the
rest of the analyzed accessions of D. rufescens and Far Eastern accessions earlier referred
to D. humilis based on their seemingly inclined capsules (yet all sporulating collections
from that area were collected with immature sporophytes, which prevented the assess-
ment of a basal membrane height). Additionally, the accession FDt119 from plants mor-
phologically approaching D. humilis collected in Sakhalin Island was found to be molecu-
larly distinct, in a sister position to the maximally supported D. rufescens clade. The topol-
ogy between the maximally supported D. rufescens + humilis clade, D. crispa + subulata
clade, and the crown clade of Dicranales has not been resolved, even with our deeper
molecular sampling, yet the analysis of trnK data alone (Supplementary Figure S2) yielded
an unsupported clade containing D. rufescens + humilis plus D. crispa + subulata lineages.
Stronger support (BI PP 1 but without support from ML) was found for the clade contain-
ing these two lineages and the clade containing the core Dicranales. The ambiguous affin-
ities of these two lineages might partly result from the ambiguous affinities of the Archi-
diaceae + Leucobryaceae clade, which was resolved in the sister position to the clade con-
taining Grimmiales based on the LTRN data (now with BS 94/PP 1 support), as opposed
to the unsupported (BS < 50/PP 0.79) sister position of the Archidiaceae + Leucobryaceae
clade to the rest of the Dicranales, excluding D. rufescens + humilis and D. crispa + subulata
lineages, in the analysis of K data alone. The signal from LTRN data was stronger than
that of K data, weakening the support for Archidiaceae/Leucobryaceae plus Grimmiales
clade to BS 67/PP 0.94 in the combined LTRKN analysis.

Plants 2023, 12, 1360 6 of 40
Figure 2. Sporophytes and peristome of Dicranella staphylina (from North Siberia, MW9117945,
FDt107). (A) fragment of specimen with sporophytes, (B) wet opening capsule with dehiscent an-
nulus, (C–E) dry capsules, (F) SEM image of peristome, (G) outer surface of the lower part of seg-
ment, (H) spore. Scale bars: 1 mm for (A); 0.5 mm for (B–E); 100 μm for (F); 10 μm for (G); 2 μm for
(H).
Support for the core Dicranales clade has grown substantially in both ML (BS 85–100
according to the dataset) and BI (PP 1), and similar support was obtained for the sister
relationship between Amphidium and the rest of the core Dicranales clade. The same is true
for the three larger clades within the core Dicranales, the first one including Schistostega,
Rhabdoweisiaceae, Ditrichaceae, Pottiaceae, Bruchiaceae, and a maximally supported
clade containing accessions of Rhamphidium, Symblepharis krausei, and Dicranella vaginata;
the second containing Dicranaceae, Fissidentaceae, Chrysoblastella, Bryowijkia, and Di-
cranellaceae; and the third containing Aongstroemiaceae.
The nearly maximally supported (BS 98–100/PP 1) Dicranellaceae clade does not con-
tain northern representatives of Dicranella s.l., except for the genus in its amended delim-
itation itself, and the newly analyzed accession of Dicranella polii from Madagascar ap-
peared in a poorly supported clade with two accessions of the genus Garckea, which itself
appeared nested within a maximally supported clade containing accessions of Microcam-
pylopus, Campylopodium, and Leptotrichella flaccidula. This clade is resolved in sister posi-
tion to the clade containing Aongstroemia filiformis s.lat. accessions. The Madagascan and

Plants 2023, 12, 1360 7 of 40
Reunion accessions of that species appear molecularly distinct from the Neotropical ac-
cessions, which earlier were recognized as A. jamaicensis Müll. Hal. Within the Dicranella
s.str. clade, accessions of D. cerviculata formed a clade sister to the remaining accessions
of the genus. Within the latter, two molecularly identical accessions of D. heteromalla/Cam-
pylopus pyriformis-like plants from Southern Kuril Islands are separated in a maximally
supported clade and are described below as Dicranella thermalis. The maximally supported
sister clade to the D. thermalis clade consists of (1) the nearly maximally supported clade
containing accessions of D. curvipes and (2) the unsupported clade containing the acces-
sions of D. heteromalla. Within this clade, two smaller clades can be recognized, one with
high support containing both European and non-European plants referable to this species,
and the other unsupported clade containing only accessions from the Russian Far East,
which might in the future receive formal status upon a detailed account of their molecular
and morphological variability.
Within the now maximally (except for ML BS 87 in the LTRN-based tree) supported
Aongstroemiaceae sensu [10], the sister group relationship between Dicranella varia s.lat.
+ D. howei + D. pacifica clade and the rest of the taxa was confirmed, yet the clade contain-
ing Aongstroemiaceae without the D. varia group now only has weak support (BS 63–78
only from K data and PP 0.67–0.97 according to dataset and indel scoring), with respect to
the inclusion of the newly analyzed basalmost lineage containing the plants described be-
low as a new genus, Protoangstroemia. Within the D. varia group, four maximally or nearly
so supported lineages could be recognized. Apart from D. varia and D. howei, the newly
analyzed D. pacifica appeared sister to D. varia + howei clade, and four accessions, contain-
ing the RF42, which earlier was assigned to D. varia but now is referred to Dicranella varia
var. obtusifolia Berggren raised to the species rank below, form a lineage sister to the rest
of the entire D. varia group. Within the Aongstroemiaceae s.str. clade, the basal grade con-
sists of Protoangstroemia and the maximally supported lineages of Diobelonella, Dichodon-
tium, and Neodicranella. However, the clade containing accessions of Neodicranella appears
in a different position in the analysis of LTRN and K data (cf. Figures S1 and S2), essen-
tially unresolved in the grade between Diobelonella and Dichodontium according to LTRN
data but deeply nested within Aongstroemia s.lat., sister to the Dicranella grevilleana +
Aongstroemia longipes clade according to K data. The crown clade (Aongstroemia s. lat.) con-
tains the Dicranella grevilleana + Aongstroemia longipes, Dicranella schreberiana var. robusta,
D. schreberiana var. schreberiana, Hygrodicranum bolivianum, H. herrerae, and Dicranella cam-
pylophylla + D. hookeri + Polymerodon andinus clades. Additional accessions of Aongstroemia
longipes, Dicranella grevilleana, D. schreberiana var. schreberiana, D. campylophylla, and D.
hookeri support the distinctness of Aongstroemia longipes from Dicranella grevilleana, Di-
cranella schreberiana var. robusta from var. schreberiana, and of D. campylophylla from D.
hookeri, yet a more detailed study of taxa in this group needs to be performed in the future
with respect to one isolated accession of Dicranella schreberiana s.lat. from Russia (RF40),
the similarly isolated accession of D. hookeri RF65, and the nested position of D. cam-
pylophylla TJH13 within the clade, which otherwise contained specimens referable to D.
hookeri.
3. Discussion
Our trees are largely congruent with those published by [10] and [23], yet bring more
resolution to the relationships among the haplolepidous lineages, identify affinities of the
seven previously unsampled species, and verify the previously assessed affinities using
additional accessions of previously insufficiently sampled taxa.
Dicranella staphylina. The totally orphaned position of Dicranella staphylina within
the system of haplolepidous mosses came as one the most surprising results of the phylo-
genetic reconstruction by [10]. This moss has to date been known from Europe essentially
only from its gametophytic stage, which does not have any distinct autapomorphic traits.
Several immature sporophytes have only been observed by [27]. They had yellow to or-

Plants 2023, 12, 1360 8 of 40
ange seta (speculated to be red at maturity), erect, symmetrical, smooth capsules with ir-
regular and incrassate exothecial cells and few stomata, longitudinally striate peristome
teeth bifid to the middle, and unmatured spores 15–20 µm. The character of annulus was
not mentioned. Neither of these characters is outstanding among northern Dicranella s.lat.
species. Unexpectedly, sporulating plants not matching the description of any other
known species of the genus were recently discovered in the north Siberian Putorana Plat-
eau. They had yellow setae, asymmetric furrowed capsules, bright red peristome, and
well-developed revoluble annulus (Figure 2). Surprisingly, the molecular barcoding of
these plants revealed their identity with the previously analyzed accessions of D. staph-
ylina and, indeed, the gametophytic characters matched the European material except for
plants from Khabarovsk Territory, discovered upon the subsequent herbarium revision,
which lack the characteristic rhizoidal tubers. The difference in the capsule shape as com-
pared to [27] might result from the ontogenetic stage, when young straight and smooth
capsules may also become curved and furrowed with age. With respect to the isolated
position of Dicranella staphylina, we propose a new genus and family to accommodate it.
The sister group relationship between D. staphylina and the clade containing both the
rest of the order Dicranales (including Archidiaceae and Leucobryaceae) and the species
currently recognized within the order Grimmiales opens the question of the ordinal place-
ment of the lineage containing D. staphylina. While the resolved topology based on LTRN
and K data differs in the assessment of affinities of the Archidium + Leucobryaceae clade,
both topologies agree on the nested position of the currently recognized Grimmiales
within Dicranales, should the D. staphylina-lineage remain in Dicranales. D. staphylina has
a fairly typical dicranoid peristome with triangular, in basal and median part longitudi-
nally striolate rather than filiform teeth, split to half of their length into unequal lobes,
without a basal membrane (Figure 2), which clearly fits the description of the dicranoid
peristome type by [28]. This favors the idea of including this lineage in the delimitation of
Dicranales, rather than establishing an isolated new order to accommodate it, suggesting
that the dicranoid peristome is the plesiomorphic character state for the whole large line-
age, from which the more derived peristomes in Grimmiales and Pottiaceae evolved (see
also [9] for a discussion of the secondarily modified peristomes in, e.g., Glyphomitrium
Brid. or Pseudoblindia Fedosov, M. Stech & Ignatov of Rhabdoweisiaceae). The idea of a
broad Dicranales, with the currently recognized Grimmiales being lowered to the rank of
suborder, has further support from the absence of any derived morphological trait that is
typical for the earlier diverging lineages, such as Catoscopium Brid., Distichium Bruch &
Schimp., Bryoxiphium Mitt., or Pseudoditrichum Steere & Z. Iwats.
Dicranella rufescens and D. humilis. These two species share the red color of their
stems [29], rather sparsely foliated stems with leaves hardly homomallous or secund,
plane leaf margins (which however can be narrowly recurved on one side in D. humilis),
and a weakly differentiated costa, especially in D. rufescens. A unique character of D. ru-
fescens among other ex-Dicranella species is the high basal membrane (up to 10 rows),
while the membrane of D. humilis does not extend four rows; basal membranes extending
three rows are nevertheless rare in all other species except D. varia. Both species markedly
differ in their capsule shape (characteristically straight and symmetric in D. rufescens,
while inclined, slightly curved, and asymmetric in D. humilis). It was therefore important
to confirm that D. humilis is indeed closely related to D. rufescens, which we accomplished.
All Asian specimens referred to D. humilis on the available morphological grounds ap-
peared in the D. rufescens clade. Thus, although our study confirmed the species status of
D. humilis, further morphological study of additional Asian specimens is needed to clarify
the differentiation of D. humilis and D. rufescens. The isolated position of the clade pre-
cludes any other taxonomic solution except for establishing a new genus for the two spe-
cies of the lineage, with the familial placement being somewhat ambiguous. The very
weak clustering with the Dicranella subulata + crispa based in trnK data (Figure S2) might
favor creating a family harboring both these lineages; however, the total evidence from all

Plants 2023, 12, 1360 9 of 40
studied regions (Figure 1) does not support this solution and favors the creation of a sep-
arate family for this monogeneric lineage. This is supported by the salient morphological
differences between the lineages (absent versus well-developed revoluble annulus, basal
membrane 3–10 versus 1–3 rows, sparse leaves, never clasping and shouldered versus
leaves dense and contiguous, at least perichaetial leaves clasping) in the absence of other
nonhomoplasic common characters.
Dicranella subulata and D. crispa. The morphological synapomorphies of this line-
age were discussed at length by [10]. While the affinities with the preceding lineage have
not been convincingly resolved, the same set of arguments can be used for segregating the
two known representatives of this lineage to a separate genus and family (see Taxonomic
treatment). The previous names adopted for Dicranella crispa and D. subulata include either
names that are in use for other distinct genera, or the illegitimate genus names Dicranodon
Béhéré and Leptotrichum Hampe ex Müll. Hal. Similarly, the possibility of raising Di-
cranella sect. Pseudodicranella Nyholm to the genus level is prevented by the name being
invalid with respect to a missing Latin description and is illegitimate, as it includes the
conserved type of Dicranella, D. heteromalla. Therefore, we propose to erect a new genus
and family name for this group in the Taxonomy section.
Dicranellaceae. In agreement with [10], we concur with the proposal of reducing the
delimitation of Dicranellaceae to only include members of the Dicranella heteromalla group,
Microcampylopus/Leptotrichella/Garckea/Campylopodium polytomy, Aongstroemia filiformis
s.lat., Eccremidium, and Cladophascum. With its remarkably distinct morphology [30], Bry-
owijkia, albeit robustly supported molecularly as a sister group to the above-specified as-
semblage, should remain separate at the family rank, as proposed by [31]. The question of
the inclusion of Trichodontium falcatum (R. Br. bis) Fife remains open. The species was
merged with Kiaeria pumila (Mitt.) Ochyra by [32], which was resolved as a member of
Arctoa within Rhabdoweisiaceae [9]. However, the only available Trichodontium GenBank
accessions AF435304/AF435353 from the specimen Streimann 51155 appeared in the clade
with Leptotrichella flaccidula and Campylopodium medium in the analysis of [10], which sug-
gests the possibility of an incorrect identification in one of the treatments, and the matter
needs to be revisited in the future.
The maximally supported clade containing accessions of Aongstroemia filiformis s.lat.
sister to the clade of other tropical Dicranellaceae also leaves us with no other option than
to establish a new genus to accommodate it, as the species has never been included in
genera other than Aongstroemia, Dicranella, Dicranum, and Thysanomitrion, which cannot
be used for this purpose. This solution is put into effect later in the text. The geographically
meaningful pattern of molecular variability, as supported by the analysis of an additional
Aongstroemia filiformis specimen from Madagascar, seems to support the resurrection of A.
jamaicensis from the synonymy of A. filiformis, but this task requires additional sampling
and morphological study.
The well-supported tropical Dicranellaceae clade, consisting of analyzed accessions
of Aongstroemia filiformis s.lat., Microcampylopus, Leptotrichella, Garckea, and Campylopodium
also contains a single analyzed specimen of Dicranella polii. Its closest affinities were re-
vealed to be with the previously analyzed Garckea species, with which it forms a clade
moderately supported from ML and not supported from BI (BS 77–81, PP 0.86–0.88),
nested within the well-supported clade containing the tropical Dicranellaceae, except for
Aongstroemia filiformis s.lat. While the combination of D. polii under Garckea would make
sense from a nomenclatural point of view, as the latter appears to be the oldest available
generic name in this clade (Leptotrichella incl. the younger Microdus, Microcampylopus), the
sporophytic characters currently used for delimitation between Garckea, Leptotrichella
(generally considered synonymous to Dicranella [20,33,34], and Microcampylopus do not
match the revealed phylogenetic affinities, and hence we prefer to postpone this taxo-
nomic decision, pending a deeper sampling in this lineage. This brings, however, another

Plants 2023, 12, 1360 10 of 40
piece of evidence that the tropical species referred previously to Dicranella s.lat., Lepto-
trichella, and Microdus might belong to this lineage, or to the lineage containing Rhamphid-
ium species (see below for a discussion of Dicranella vaginata).
The affinities within Dicranella s.str. support the description of a new species, as re-
alized below, and the continued recognition of D. curvipes from D. heteromalla at the spe-
cific rank, as suggested by [35], with the molecular support presented in a more limited
dataset by [10]. Our additional data support the recognition of Dicranella curvipes as a sep-
arate entity, while further documenting the molecular variability within the lineage. Our
review of gametophytic morphology revealed that, in some of Dicranella curvipes speci-
mens, the leaves tend to be homomallous vs. mostly falcate secund in D. heteromalla; the
costa is narrower (less than 1/5 of the leaf base width), well-delimited from leaf lamina (a
unique trait in Dicranella s.str.), and unistratose throughout (vs. wider costa weakly de-
limited from leaf lamina, which is partly to nearly entirely bistratose distally in D. heter-
omalla); the cells in the basal leaf portion are narrowly rectangular and moderately thick-
walled vs. short-rectangular to subquadrate, thin-walled in D. heteromalla). In addition,
most specimens of D. curvipes have leaves with rather distinct shoulders. It needs to be
acknowledged, though, that some specimens of D. curvipes (such as Kučera 21379, 21778)
from the Russian Far East have other combinations of these characters and could not be
identified without mature sporophytes. Moreover, there appears to be an internal differ-
entiation of the clade consisting of plants currently assigned to D. heteromalla s.str., with
the specimens from the Russian Far East (RF47, 49, FDt35, Kučera 21639) and one from the
eastern United States (Goffinet 8162) showing several distinct molecular synapomorphies
at the level of one-base substitutions. Although this clade is only weakly supported on the
tree, with respect to the unequal sequenced regions in the studied accessions and ambig-
uous reads at several points, the lineage is probably molecularly distinct. The most di-
verged lineage in molecular terms is, however, the one harboring two accessions of plants
collected on Iturup Island and originally identified as Campylopus pyriformis. Despite the
few morphological traits differentiating these plants from D. heteromalla, the plants are
described below as a morphologically semicryptic species; nevertheless, they are distinct
with respect to their rate of molecular differentiation.
Aongstroemiaceae. The common characters of Aongstroemiaceae and features
which differentiate the Dicranella varia group as the most alien element in Aongstroemi-
aceae were discussed in detail by [10]. Both morphology and molecular support for the
clade containing D. varia, D. varia var. obtusifolia, D. howei, and D. pacifica require the ge-
neric rank to be used for this clade. In theory, the name Anisothecium could be applied to
it, as Anisothecium varium is one of the six species cited in the protologue. However, we
believe that this would be the least appropriate option for the typification of the genus, as
[26] proposed this name in his “Musci Austro-Americani …” for a group of predomi-
nantly South American species, which mostly share vaginate or semivaginate leaves with
distinctly widened leaf bases, while Dicranella varia, which was included based on a single
specimen from Cuba, forms a distinctly discordant element in his circumscription of the
genus, as was also emphasized in his key to the species. Therefore, we believe that Ani-
sothecium is much more appropriately typified with one of the predominantly South
American species with expanded leaf bases, as done below, and we propose a new genus
name for the clade of Dicranella varia and closely related species.
Previous analysis [10] suggested that Dicranella varia was paraphyletic with respect
to a specimen (RF42) from northern Siberia. Extended sampling of Asian material that was
supposed to represent D. varia resulted in both plants being molecularly identical or
closely related to specimens from Europe and plants identical to the previously studied
RF42. The provenance of the latter specimens mostly included northern Siberia, while the
lineage containing European plants included specimens collected throughout boreal Asia.
Morphological examination of the north Siberian plants and comparison to D. varia s.str.,
as represented by both European specimens (the Central European lectotype from Leip-
zig, Germany, was reviewed by [36] and a specimen from the southern Siberia and the

Plants 2023, 12, 1360 11 of 40
southern part of Russian Far East), confirmed the morphological differences between the
two groups. It appeared that plants similar to the analyzed north Siberian ones had al-
ready been described. Lindberg and Arnell [37], who proceeded extensive bryophyte col-
lections from the Russian Arctic, described Anisothecium rubrum var. obtusiusculum based
on the plants from the lower course of the Yenisey River. Their description matches our
plants well. They also mentioned that a similar taxon, Dicranella varia var. obtusifolia Berg-
gren occurs in Svalbard and indeed provided a description that seems to match morpho-
logically both the variety later described by [37] and the plants that we collected in north-
ern Siberia. The type material held in MO (MO-407808, accession 2226886) shows a good
match with the north Asian plants analyzed by us. Hence, we raise the variety earliest
described by Berggren to the species rank in the newly established genus, as effected be-
low in the Taxonomy section. The previously unsampled NW American endemic Di-
cranella pacifica W.R. Schofield, which shares with D. varia multiple characters including
recurved leaf margins, inclined asymmetric capsules, and absent annulus [20,38], was con-
firmed as another member of this lineage. While the specific status of D. howei, which
morphologically sometimes seems indistinct from D. varia [20], now appears unequivocal
with respect to the resolved identity of D. varia var. obtusifolia, the elaboration of morpho-
logical differences remains the task for a future dedicated study with more numerous mo-
lecularly barcoded specimens.
The core Aongstroemiaceae clade contains two basal lineages, which were not sam-
pled by [10]. The basalmost lineage is represented by a single collection of a plant from
Sakhalin Island with a unique combination of the otherwise typical Aongstroemiaceae
characters, including shouldered leaves; elongate laminal cells; a single costal stereid band
and lack of guide cells; reddish setae; and short, dark, curved, smooth, or slightly fur-
rowed exannulate capsules. On the contrary, the upper leaf lamina is bistratose to tristra-
tose, a character that only occurs in some representatives of the family. This plant is there-
fore described below as a new monospecific genus. The second previously unsampled
lineage is the likewise monotypic Neodicranella, whose affinities had not been well re-
solved in its protologue [23]. We were not able to convincingly assess its affinities, even
now, due to the conflict in the resolved topologies between the LTRN and K datasets.
Although the “dicranelloid” rather than “dichodontioid” habit would favor the affinities
as assessed through analysis of the trnK data, we refrain, however, from merging Neodi-
cranella with Aongstroemia at this point.
The crown clade of core Aongstroemiaceae contains the species recognized to date,
aside from the above-mentioned conflicting position of Neodicranella, in at least four gen-
era: Aongstroemia (type species A. longipes), Hygrodicranum (type species H. falklandicum
Cardot, not analyzed), Polymerodon (monotypic), and Dicranella species with expanded,
mostly vaginate leaf bases, which earlier were often assigned to Anisothecium (the ana-
lyzed Dicranella campylophylla is among the six Anisothecium species eligible for the type of
the genus). The high molecular support for this clade, as well as the suite of morphological
characters common in the analyzed taxa of this clade, strongly support the idea of recog-
nizing this clade as one genus, for which the oldest available name is Aongstroemia (see
below in the Taxonomy section). Within the genus, we were able to additionally analyze
four specimens of Dicranella schreberiana s.str. from previously unsampled parts of Europe
(Czech Republic, Greece, European Russia), which confirmed the distinctness from D.
schreberiana var. robusta. Moreover, we morphologically revised the type specimen of
Cynodontium canadense Mitt., which proved to be identical to D. schreberiana var. robusta.
Having priority at species rank, Cynodontium canadense is combined under Aongstroemia
later in the Taxonomy section, leaving however open the question of the specimen RF40,
with the morphology rather suggesting D. schreberiana s.str., which was however found
to be isolated from the clades representing both recognized varieties and did not form a
monophylum with either of them. Additional analyzed specimens of D. campylophylla (in-
cluding the specimens labelled as D. cardotii (R. Br. bis) Dixon, considered synonymous
by [39] and subsequent authors) and D. hookeri brought more certainty to the taxonomic

Plants 2023, 12, 1360 12 of 40
evaluation of these taxa. They were found to be closely related, yet probably distinct spe-
cies, which however may be at times difficult to separate morphologically, as the accession
D. campylophylla TJH13, which was unavailable for our study, was resolved in the poly-
tomy formed by the accessions of D. hookeri, together with an accession labelled as Polym-
erodon andinus, which was downloaded from GenBank. The pattern is further complicated
by the accession RF65, identified also as D. hookeri, which however is molecularly clearly
distinct from all other accessions identified as this species, as well as from D. cam-
pylophylla. The type of Hygrodicranum, H. falklandicum Cardot remains unsampled, and
hence the generic status of Hygrodicranum remains to be assessed. Based on the two ana-
lyzed accessions of H. bolivianum and one of H. herrerae, it appears rather safe to infer that
both species are very closely related to D. campylophylla, yet possibly specifically distinct,
although the protologue and illustrations of Hygrodicranum herrerae in [40] do not provide
information that would distinguish this species from the descriptions and illustrations of
D. campylophylla available in the literature [39]. The sequenced specimens IPG20 (H. herre-
rae), as well as TJH04 and TJH13 (D. campylophylla and D. campylophylla/hookeri) from
Chile, are nearly identical, with bistratose leaf lamina and dorsally mamillose cells char-
acteristic for both taxa. The specimen of Hygrodicranum bolivianum (Buck 39497), as well as
the additionally studied Chilean specimen (Larraín 43529), matches the species description
[41], which resembles some closely related Dicranella (Aongstroemia) species. Conse-
quently, we propose to combine both H. bolivianum and H. herrerae in Aongstroemia and
expect that H. falklandicum might be resolved in this clade as well, but until the species is
analyzed, we prefer not to put this taxonomic change into effect. The same applies to the
assessment of Polymerodon andinus (rps4 and nad5 sequences obtained from specimen M.
Lewis 87608, 7/87 (DUKE) and Eucamptodontopsis pilifera (Mitt.) Broth. (nad5 sequence ob-
tained from specimen S.R. Hill 27912, 2/97 (DUKE)). If the sequences indeed correspond
to these taxa, then they should be considered conspecific with D. hookeri, but in the absence
of type studies and a more representative selection of analyzed material, such a proposal
is premature, as further corroborated by the affinities of GenBank sequences of Eucamp-
todontopsis brittoniae (E.B. Bartram) B.H. Allen (AF435285, AF435328), which appear to be
closely related to Holomitrium species based on BLAST searches.
The caution with the taxonomic evaluation of this group of taxa can be illustrated by
the example of Dicranella vaginata. This species was considered to be closely related to the
group of South Hemispheric Dicranella species recognized as Anisothecium by [26] in the
protologue of the genus. However, three Chilean accessions analyzed by us were found
to be resolved in the maximally supported clade containing two Rhamphidium species and
Symblepharis krausei (Lorentz) Ochyra & Matteri. Indeed, all species have the vaginate leaf
base, elongate basal leaf cells with porose longitudinal cell walls, subquadrate upper leaf
cells, mostly unistratose upper leaf lamina, and rather short erect or inclined, nearly sym-
metric capsule with markedly spiculose, deeply divided peristome teeth, different from
the typical dicranoid, i.e., not spiculose, and less divided peristome shared by Aongstroe-
mia species. Recent morphological studies found Dicranella vaginata very similar to
Aongstroemia gayana [42,43], which further indicates the necessity of a modern polyphasic
reassessment of the lineage containing the type of Rhamphidium.
Consequently, we propose to typify the genus Anisothecium with A. campylophyllum,
which in our opinion best preserves Mitten’s original idea to include in it mostly South
American representatives of the then recognized broad genus Aongstroemia with broad-
ened, mostly clasping leaf bases and dicranoid affinities. Neither A. varium nor A. vagina-
tum (see above) seem to qualify better for this purpose. The identity of A. jamesonii Mitt.
is currently ambiguous, as it was considered synonymous either to A. vaginatum [44] or to
Dicranella hookeri [45], which were not found to be closely related by us, and hence a new
investigation of the type is necessary in light of this finding. Similarly, we have no molec-
ular data for the remaining South American species of the original Anisothecium, A. convo-
lutum (Hampe) Mitt. and A. planinervium (Taylor) Mitt.

Plants 2023, 12, 1360 13 of 40
The polyphyly of Aongstroemia, Dicranella, and Ditrichum demonstrated by the anal-
yses in [8,10] and this study is yet another example of homoplasy of morphological char-
acters that were considered taxonomically relevant in earlier classifications. The superfi-
cially similar small pioneer mosses that are adapted to opportunistic life strategies some-
times occupy a remarkably isolated phylogenetic position among the basal lineages of
Dicranidae. They share with most other protohaplolepideous lineages the broad, typically
circumholarctic ranges, usually associated with humid climates, suggesting that the early
diversification of Dicranidae was associated with cool to mild conditions, and therefore
might be underestimated in temperate and subarctic, and by analogy possibly also in sub-
antarctic, areas. The early radiation might not have been followed by excessive diversifi-
cation according to our current knowledge, but the pioneer strategy of their representa-
tives might have allowed them to survive until the present.
In contrast, the later diverging lineages of (mostly) opportunistic pioneer mosses
(Aongstroemiaceae s.str., Dicranellaceae s.str., Ditrichaceae s.str. and some groups of Pot-
tiaceae) are remarkably more diverse, in terms of species numbers, morphologically, and
ecologically, often occupying xeric environments (such as several groups of Pottiaceae)
and displaying multiple transitions to annual life strategies. Although only a limited num-
ber of Aongstroemia and especially Dicranella species from outside the Holarctic have been
studied, the preliminary rps4-based phylogenetic analysis of Brazilian Dicranella s.lat. spe-
cies indicates that the studied Neotropical Dicranella species all belong to the lineage of
Dicranellaceae [46].
Our results demonstrate the unexpectedly underestimated diversity of northern tem-
perate and subarctic pioneer mosses with dicranelloid habit and the resulting limitations
of the currently used floras, especially in North Asia. In addition, northern Asia is an area
of higher molecular diversity of Dicranella s.l. species, while European accessions are typ-
ically uniform in sequences, which might indicate the role of northeastern Asia as a source
of diversity in these lineages worldwide.
The stunning extent of convergence in the available morphological traits within the
studied genera underlines the need for obtaining molecular data for the representatives
of the as yet unevaluated taxa and also the revision of types for existing names. Given the
number of poorly known taxa ( >600 names in Dicranella and >260 in Aongstroemia, [22]),
such a project would require the efforts of the whole bryological community.
4. Taxonomy
Rhizogemmaceae Bonfim Santos, Siebel & Fedosov, fam. nov.–Type: Rhizogemma
Bonfim Santos, Siebel & Fedosov
Diagnosis: This family differs from the other families of haplolepideous mosses in
possessing the following combination of characters: plants small to medium-sized; stems
with central strand; leaves with widened leaf bases abruptly narrowed to short subulate
leaf tips; leaf margins recurved; costae with central stereid band, dorsal and ventral epi-
dermis, without guide cells; laminal cells elongate, smooth; sexual condition dioicous; se-
tae yellow; capsules asymmetric, furrowed; peristome dicranoid, bright-red; annulus
revoluble.
The family is currently considered monogeneric.
Rhizogemma Bonfim Santos, Siebel & Fedosov, gen. nov.–Type: Rhizogemma staph-
ylina (H. Whitehouse) Bonfim Santos, Siebel & Fedosov (Figures 2 and 3).

Plants 2023, 12, 1360 14 of 40
Figure 3. Line drawings of gametophyte and sporophyte of Siberian plants of Rhizogemma staphylina
(from Russia: Krasnoyarsk Territory, Putorana Plateau, MW9117945, isolate FDt107): (A) fertile
plant, wet, (B) fertile plant, dry, (C–E,O) capsules, dry, (F) exothecial cells, (G–J) leaf transverse
sections, (K,P) perichaetial leaves, (L,M,Q,R,T,U,V) stem leaves, (N,W) upper-leaf cells, (S,X) mid-
leaf cells, (Y,BB) basal leaf cells, (Z,AA) gemmae. Scale bars: 2 mm for (A,B); 0.5 mm for (C–E,O); 1
mm for (K–M,P–R,T,U,V); 100 μm for (F–J,N,S,W–BB).
Diagnosis: The single species segregated into the newly established genus differs
from other dicranelloid mosses in possessing non-vaginate leaf bases and rather shortly
subulate leaf acumina, recurved leaf margins, costae with single central stereid band, leaf
lamina unistratose or bistratose along upper margins, yellow setae, yellow-purplish to
brownish, asymmetric, furrowed capsules, revoluble annulus, and rhizoidal gemmae ir-
regular in shape, composed of bulging cells.
Etymology: The name (composed of the Greek ῥίζα, root, and Latin gemma, gem)
refers to the characteristic rhizoidal tubers (commonly also referred to as gemmae) of the
only currently known species of the genus.
Description: Plants bright green, lacking red pigmentation. Stems about 5 mm, form-
ing rather dense tufts, with a central strand. Leaves up to 1 mm long, lanceolate, erect-
spreading to spreading, not secund; margins plane or recurved only at base or nearly
throughout, smooth or denticulate distally, partly bistratose distally; costae percurrent to
short excurrent, in transverse section with differentiated dorsal and ventral epidermis and
single stereid band; leaf lamina unistratose, cells rectangular to elongate-rectangular,
bulging in transverse sections, smooth. Rhizoidal tubers constantly present, in young

Plants 2023, 12, 1360 15 of 40
stage red, turning dark brown, irregularly shaped with protruding cells, 3−4 cells long
and 1−3 cells wide. Perichaetial leaves differentiated, larger than lower leaves, from
broadly sheathing base rather abruptly narrowed into squarrose or flexuose apex. Setae
yellowish, straight; capsules incurved, longitudinally furrowed, without or weak stru-
mae, yellow to purplish along ribs and around the mouth. Exothecial cells irregular in
shape to rectangular, with evenly incrassate walls. Annulus well differentiated, composed
of one row of large hyaline thick-walled cells, revoluble. Operculum long rostrate. Peri-
stome bright red, teeth unequally split to half of their length, longitudinally striolate prox-
imally, papillose distally. Calyptrae cucullate.
The genus is currently considered monotypic.
Rhizogemma staphylina (H. Whitehouse) Bonfim Santos, Siebel & Fedosov, comb.
nov. ≡ Dicranella staphylina H. Whitehouse in Trans. Brit. Bryol. Soc. 5: 757. f. 2-3a. 1969.–
Type: United Kingdom, E. Norfolk (v.-c. 27), Pockthorpe, near Lyng, arable field, Sept.
1968, H.L.K. Whitehouse s.n. (Holotype: CGE) ≡ Anisothecium staphylinum (H. Whitehouse)
Sipman, Rubers & Riemann in Lindbergia 1: 217. 1972.
Ruficaulaceae Bonfim Santos & Fedosov, fam. nov.–Type: Ruficaulis Bonfim Santos
& Fedosov
Diagnosis: This family differs from the other families of haplolepideous mosses in
possessing the following combination of characters: small plant size; mature stems red-
dish-brown; costae with only dorsal stereid band; leaf lamina unistratose or with bistra-
tose margin, composed of elongate to linear cells; sexual condition dioicous; red setae;
peristome dicranoid, with rather weak ornamentation, well-developed basal membrane
and weakly developed annulus.
The family is currently considered monogeneric.
Ruficaulis Bonfim Santos & Fedosov gen. nov.–Type: Ruficaulis rufescens (With.) Bon-
fim Santos & Fedosov
Etymology: The generic name originates from the Latin caulis, stem, and the prefix
rufi- (from Latin rufus, red), referring to the characteristic reddish-brown coloration of the
stem.
Diagnosis: species combined in the newly established genus differ from other di-
cranelloid mosses in possessing reddish-brown mature stems; leaves from narrow-trian-
gular base gradually narrowed to a subulate acumina; costae with single stereid band; leaf
lamina unistratose or with bistratose margins, composed of elongate to linear cells; red
setae; peristome with mostly high basal membrane and weakly developed annulus.
Description: Plants very small, in loose reddish-brownish tufts. Stems with central
strand. Well-developed parts of stems reddish-brown and rhizoids with vinaceous pig-
mentation. Leaves up to 2 mm long, fuscous, weakly secund; margins plane throughout,
denticulate at apex; costa rather weak, percurrent, sharply delimited from leaf lamina,
with compact stereid band, ventral and dorsal epidermis, or with weakly developed dor-
sal band, composed of substereids and guide cells covered by ventral epidermis. Tubers
consisting of one row of (1−)2−3(−6) much enlarged cells, pale reddish to wine-red.
Perichaetial leaves similar to upper leaves. Setae reddish. Capsules erect to inclined, sym-
metric or curved, smooth or slightly furrowed. Exothecial cells short rectangular, in lon-
gitudinal rows. Annulus weakly differentiated, not revoluble. Peristome dicranoid, with
high basal membrane.
Accepted species:
Ruficaulis rufescens (With.) Bonfim Santos & Fedosov, comb. nov. ≡ Bryum rufescens
With. in Syst. Arr. Brit. Pl. (ed. 4) 3: 801. 1801–Type: “ad ripas rivulorum lutosas, in Scotia”
≡ Dicranum rufescens (With.) Turner in Muscol. Hibern. Spic. 66. 1804 ≡ Dicranum varium
var. rufescens (With.) Röhl. in Deutschl. Fl. (ed. 2), Kryptog. Gew. 3: 71. 1813 ≡ Dicranodon
varium var. rufescens (With.) Béhéré in Muscol. Rothom. 29. 1826 ≡ Dicranum crispum var.
rufescens (With.) Hampe in Flora 20: 283. 1837 ≡ Aongstroemia rufescens (With.) Müll. Hal.
in Syn. Musc. Frond. 1: 436. 1848 ≡ Dicranella rufescens (With.) Schimp. in Coroll. Bryol.
Eur. 13: 1856 ≡ Anisothecium rufescens (With.) Lindb. in Musci Scand. 26. 1879.

Plants 2023, 12, 1360 16 of 40
Ruficaulis humilis (R. Ruthe) Jan Kučera & Fedosov, comb. nov. ≡ Dicranella humilis
R. Ruthe, Hedwigia 12: 147. 1873.–Type: [Germany] In locis paucis prope Bärwalde Ne-
omarchicae. ≡ Anisothecium humile (R. Ruthe) Lindb., Meddeland. Soc. Fauna Fl. Fenn. 14:
74. 1887 ≡ Aongstroemia humilis (R. Ruthe) Müll. Hal., Gen. Musc. Frond. 323. 1900. ≡ Di-
cranella rufescens subsp. humilis (R. Ruthe) Kindb., Eur. N. Amer. Bryin. 2: 209. 1897.
Dicranellopsidaceae Bonfim Santos, Siebel & Fedosov, fam. nov.–Type: Dicranellop-
sis Bonfim Santos, Siebel & Fedosov
Diagnosis: This family differs from the other families of haplolepideous mosses in
possessing the following combination of characters: plants small to medium-sized; stems
with central strand; leaves with widened to vaginate leaf bases, abruptly narrowed to su-
bulate leaf acumina; leaf margins plane; costae with two stereid bands and guide cells;
leaf lamina bistratose distally; laminal cells elongate, smooth; sexual condition dioicous;
red setae; ribbed capsules; Dicranoid peristome and revoluble annulus.
The family is currently considered monogeneric.
Dicranellopsis Bonfim Santos, Siebel & Fedosov, gen. nov.–Type: Dicranellopsis subu-
lata (Hedw.) Bonfim Santos, Siebel & Fedosov
Etymology: The generic name originates from Dicranella, the genus to which the spe-
cies of Dicranellopsis had been assigned previously, and the suffix -opsis (from Greek ὄψις,
meaning aspect, appearance, sight), referring to the morphological similarity between the
genera.
Diagnosis: Species combined in the newly established genus differ from other di-
cranelloid mosses in possessing a combination of widened to vaginate leaf bases and su-
bulate leaf tips, plane leaf margins, costae with two stereid bands and guide cells, bistra-
tose upper leaf lamina, red setae, ribbed capsules and revoluble annulus.
Description: Plants yellowish green to dark green, lacking red pigmentation. Stems
with central strand. Leaves with oblong bases abruptly tapering into a long, channeled,
subulate acumina, upper stem leaves sheathing, erect to squarrose-flexuose, patent or se-
cund, crispate or not when dry; margins entire or very slightly denticulate at leaf tip,
plane, unistratose; costae percurrent to short excurrent, sharply delimited from leaf lam-
ina, with dorsal and ventral epidermis, guide cells and dorsal and ventral or only dorsal
stereid band; distal leaf lamina bistratose, median leaf cells linear. Rhizoidal tubers, when
present, dark brown, irregularly shaped without protruding cells, curved. Perichaetial
leaves resemble upper stem leaves. Capsules erect to slightly inclined, symmetric or dis-
tinctly curved, not strumose, strongly longitudinally ribbed, with more or less differenti-
ated exothecial bands and quadrate to short rectangular, rather thin-walled cells between
them. Annulus differentiated in 2‒3 rows of widened cells, revoluble. Peristome dicra-
noid.
Accepted species:
Dicranellopsis crispa (Hedw.) Bonfim Santos, Siebel & Fedosov, comb. nov. ≡ Di-
cranum crispum Hedw. in Sp. Musc. Frond. 132. 1801–Lectotype: Sweden, J.F. Ehrhart s.n.
(G, barcode G00040017, [36]: Figure 1B–D; [47]: Figure 51) ≡ Aongstroemia crispa (Hedw.)
Müll. Hal. in Syn. Musc. Frond. 1: 439. 1848 ≡ Dicranella crispa (Hedw.) Schimp. in Coroll.
Bryol. Eur. 13. 1856 ≡ Leptotrichum crispum (Hedw.) Mitt. in J. Proc. Linn. Soc., Bot., Sup-
plementary 1: 158. 1859 ≡ Cynodontium crispum (Hedw.) Mitt. in J. Proc. Linn. Soc., Bot. 8:
15. 1864 ≡ Anisothecium crispum (Hedw.) C.E.O. Jensen in Skand. Bladmossfl. 314. 1939,
nom. illeg., non Lindb. in Utkast Eur. Bladmoss. 33. 1878.
Dicranellopsis subulata (Hedw.) Bonfim Santos, Siebel & Fedosov, comb. nov. ≡ Di-
cranum subulatum Hedw. in Sp. Musc. Frond. 128. T. 34. f. 1-5. 1801–Lectotype: Sweden,
Swartz s.n. (G, Hb. Hedwig-Schwägrichen, barcode G00040102, [36]: Figure 5A–C, [47]:
Figure 61) ≡ Dicranodon subulatum (Hedw.) Béhéré in Muscol. Rothom. 29. 1826 ≡ Dicranum
heteromallum var. subulatum (Hedw.) Wallr. in Fl. Crypt. Germ. 1: 160. 1831 nom. illeg. ≡
Aongstroemia subulata (Hedw.) Mül. Hal. in Syn. Musc. Frond. 1: 433. 1848 ≡ Dicranella su-
bulata (Hedw.) Schimp. in Coroll. Bryol. Eur. 13: 1856 ≡ Leptotrichum subulatum (Hedw.)

Plants 2023, 12, 1360 17 of 40
Mitt., in J. Proc. Linn. Soc., Bot., Supplementary 1: 9. 1859 ≡ Cynodontium subulatum
(Hedw.) Mitt. in J. Proc. Linn. Soc., Bot. 8: 15. 1864.
Dicranellaceae Stech in Nova Hedwigia 86: 14. 2008–Type: Dicranella (Müll. Hal.)
Schimp.
Accepted genera: Campylopodium (Müll. Hal.) Besch. (only C. medium studied), Clad-
ophascum Dixon, Dicranella (Müll. Hal.) Schimp. Eccremidium Wilson (only E. floridanum
studied), Garckea Müll. Hal., Leptotrichella (Müll. Hal.) Lindb. (only L. flaccidula studied),
Microcampylopus (Müll. Hal.) Fleisch., Bryopalisotia Bonfim Santos & Fedosov.
Tentatively included genus (pending molecular confirmation): Bryotestua Thér. & P.
de la Varde. As for Trichodontium (Dixon) Fife, see the Discussion.
Description: Plants small to medium-sized, growing in turfs. Acrocarpous or clado-
carpous. Central strand present. Leaves appressed or erect-spreading, often flexuose or
falcate-secund, narrowly lanceolate, often subulate. Lamina cells rectangular, smooth, not
porose. Alar cells not differentiated. Costa single, strong, (sub-)percurrent to (long) excur-
rent, with guide cells, dorsal and ventral stereid bands and differentiated ventral and dor-
sal epidermis. Dioicous or autoicous. Seta elongate, erect, sinuose or arcuate, or short,
erect or curved. Capsule erect to horizontal or pendulous, symmetric or gibbous, occa-
sionally strumose, smooth or plicate, ovoid to short-cylindric with operculum conic to
long-rostrate, or globose with operculum dome-shaped to hemispheric with a blunt apicu-
lus. Stomata present or absent. Peristome dicranoid or absent. Spores usually papillose,
sometimes warty. Calyptra cucullate or mitrate.
The following synopsis only includes the genera where taxonomic novelties are pro-
posed.
Dicranella (Müll. Hal.) Schimp. in Coroll. Bryol. Eur. 13. 1856–Type: Dicranella heter-
omalla (Hedw.) Schimp.
Accepted species: Dicranella cerviculata (Hedw.) Schimp., D. curvipes (Lindb.) Ignatov,
D. heteromalla (Hedw.) Schimp., D. thermalis Fedosov & Ignatova (see below).
Excluded species: Dicranella campylophylla (Taylor) A. Jaeger, D. crispa (Hedw.)
Schimp., D. grevilleana (Brid.) Schimp., D. hookeri (Müll. Hal.) Cardot, D. howei Renauld &
Cardot, D. humilis Ruthe, D. pacifica W.B. Schofield, D. riparia (H. Lindb.) Mårtensson &
Nyholm, D. rufescens (With.) Schimp., D. schreberiana (Hedw.) Hilf. ex H.A. Crum & L.E.
Anderson, D. staphylina H. Whitehouse, D. subulata (Hedw.) Schimp., D. varia (Hedw.)
Schimp.
Species with uncertain placement: all other accepted species (cf. [22]), pending mor-
pho-molecular studies, including Dicranella polii Renauld & Cardot and D. vaginata
(Hook.) Cardot, for which our molecular phylogenetic data suggest placement outside
Dicranella as recognized here, but additional sampling is needed to assign the generic af-
finities, as discussed above.
Dicranella thermalis Fedosov, Ignatova & Jan Kučera, sp. nov. (Figure 4).

Plants 2023, 12, 1360 18 of 40
Figure 4. Line drawings of gametophyte of Dicranella thermalis (from: Holotype, isolate CF1-1): (A–
C) leaf transverse sections, (D,E) view of sterile plants, (F–H) leaves, (I) upper-leaf cells, (J) mid-leaf
cells, (K) basal leaf cells. Scale bars: 5 mm for (D); 2 mm for (E); 1 mm for (F–H); 100 μm for (A–C),
(I–K).
Diagnosis: The new species resembles D. heteromalla in the rather robust plant size,
non-shouldered leaves, wide costae occupying up to ½ of the leaf width and weakly de-
limited from leaf lamina, with thin-walled cells with large lumen forming ventral surface
of costa in basal leaf portion, but differs from it in having homomalous rather than falcate
secund leaves and weakly serrulate to nearly entire upper leaf margins.
Type: Russia, Sakhalin Province, Iturup Island. South-West slope of Baranskogo Vol-
cano, Goryachaya River. Fedosov & Pisarenko 19 September 2015, Mosses of the Russian
Far East Exsiccatae No. 78 (as Campylopus pyriformis). (Holotype: MW: MW9090383, Iso-
types: MHA, NSK, VGBI, MO, NY).
Etymology: The species name refers to the typical habitat of the species at the type
locality.
Description: Plants medium-sized, stems up to 3 cm, single, with well-developed
central strand, evenly foliate, tomentose in lower part. Leaves more or less appressed
when dry, spreading when wet, gently falcate-secund, 2.5 – 3.2 × 0.25 – 0.35 mm, widest
at base, from lanceolate base gradually tapering into canaliculate subulate acumen; mar-
gins plane, unistratose, weakly and bluntly toothed throughout or only in upper half, near
apex with double teeth: costae strong, occupying 1/3–1/2 of leaf base, rather indistinctly

Plants 2023, 12, 1360 19 of 40
delimited from the leaf lamina, with one row of guide cells, two stereid bands, and differ-
entiated dorsal and ventral epidermis; sometimes ventral epidermis immediately cover-
ing guide cells or guide cells forming surface of costa ventrally; leaf lamina partly or com-
pletely bistratose distally, upper leaf cells 24 – 38 × 5 – 6 µm, elongate-rectangular, smooth,
moderately thick-walled; basal leaf cells of the same length and 8–11 µm wide. Sexual
condition and sporophytes unknown.
Differentiation: We did not find more characteristics to differentiate this molecularly
distinct species from Dicranella heteromalla than those specified in the diagnosis. D. ther-
malis resembles Campylopus pyriformis (Schultz) Brid. in having wide costae, undifferenti-
ated alar regions, and thin-walled cells with wide lumen on a ventral surface in the basal
portion of leaf. In contrast to most Campylopus species, D. thermalis possesses two stereid
bands.
Ecology and distribution: The species is known from numerous collections (held
mostly in MW) on the slope of Baranskogo volcano in Iturup Island (45.07 °N, 147.98 °E),
where it grows along the hot stream banks under Sasa understory at altitudinal range of
220–280 m a.s.l. Similar non-sporulating Dicranella plants were frequently encountered in
thermal habitats of Kamchatka Peninsula and northern part of Kunashir Island, but they
were mostly not collected and therefore their identity remains uncertain.
Paratypes (the same locality, date and collectors as in the holotype): Accession num-
bers MW9073555-MW9073558, MW9007288-MW9007292, MW9073559.
Bryopalisotia Bonfim Santos & Fedosov, gen. nov.–Type: Bryopalisotia filiformis (P.
Beauv.) Bonfim Santos & Fedosov
Etymology: The name was chosen as a tribute to A.M.F.J. Palisot, Baron de Beauvois
(1752–1820), a French naturalist and author of Prodrome des cinquième et sixième familles
de l’Æthéogamie, les mousses, les lycopodes [48], in which the type species of the genus
was described as Dicranum filiforme P. Beauv.
Diagnosis: This genus differs from A. longipes and several other species of Aongstro-
emia in its traditional circumscription in its robust habit, leaves with sheathing leaf base,
abruptly narrowed into a long, subulate leaf apex, and elongate to linear, extremely thick-
walled basal leaf cells. From the genus Aongstroemia in its newly proposed circumscrip-
tion, Bryopalisotia differs in having cylindric rather than ovoid or shortly ellipsoid cap-
sules. Elongate to linear, extremely thick-walled basal leaf cells differ Bryopalisotia from A.
guayana.
The genus is presently considered monospecific, although the below-stated synon-
ymy should be revisited (see Discussion).
Bryopalisotia filiformis (P. Beauv.) Bonfim Santos & Fedosov, comb. nov. ≡ Di-
cranum filiforme P. Beauv. in Prodr. Aethéogam. 53. 1805–Type: Isle de Bourbon [=Réu-
nion], Bory s.n. ≡ Thysanomitrion filiforme (P. Beauv.) Arn. In Mém. Soc. Linn. Paris 5: 263.
1827 ≡ Aongstroemia filiformis (P. Beauv.) Wijk & Margad. in Taxon 9: 50. 1960 = Aongstroe-
mia jamaicensis Müll. Hal., Bull. Herb. Boissier 5: 554, 1897 fide [49].
Aongstroemiaceae De Not. in Atti Reale Univ. Genova 1: 30. 1869–Type: Aongstroe-
mia Bruch & Schimp.
Accepted genera: Aongstroemia Bruch & Schimp., Calcidicranella Bonfim Santos, Fedo-
sov & Jan Kučera, Dichodontium Schimp., Diobelonella Ochyra, Neodicranella Porley & Fe-
dosov, Protoaongstroemia Fedosov, Ignatova & Jan Kučera.
Tentatively included genus (pending molecular confirmation): Aongstroemiopsis M.
Fleisch. Genera tentatively moved in synonymy (see below): Hygrodicranum Cardot, Po-
lymerodon Herzog
Plants minute to medium-sized, in loose to dense turfs. Stems julaceous or not, cen-
tral strand present. Stem leaves with a broad sheathing base tapering into a blunt apex or
abruptly narrowed to a short or long acumen. Margins entire, crenulate or weakly dentic-
ulate to dentate. Leaf lamina 1–2(–3) stratose; laminal cells variable in shape, usually
smooth but mamillose or papillose in some species. Alar cells not differentiated. Costa
subpercurrent to mostly short to long excurrent, weak to strong. Asexual reproduction via

Plants 2023, 12, 1360 20 of 40
gemmae (on filamentous branches at the leaf axils) or rhizoidal tubers. Dioicous. Seta elon-
gate, straight or flexuose. Capsule variable in shape, ovoid to curved and sometimes
slightly strumose, smooth or furrowed when dry, operculate, with peristome teeth verti-
cally pitted-striolate at base. Annulus not or poorly differentiated. Operculum conic or
rostrate. Calyptra cucullate.
The following synopsis only includes genera where taxonomic novelties are pro-
posed.
Calcidicranella Bonfim Santos, Fedosov & Jan Kučera, gen. nov.–Type: Calcidi-
cranella varia (Hedw.) Bonfim Santos, Fedosov & Jan Kučera.
Etymology: The generic name originates from the generic name Dicranella, where this
species has been placed for a long time, and the prefix calci- referring to the ecological
preference for calcareous substrates in the species included in the genus.
Diagnosis: Species combined in the newly established genus differ from other di-
cranelloid mosses in possessing non-vaginate leaf bases, partly to nearly entirely recurved
leaf margins, smooth laminal cells, costae with well-differentiated stereids in one or two
bands, red setae, dark reddish-brown, asymmetric, inclined capsules, and non-revoluble
annulus, and by its ecological preference for base-rich mineral soil.
Description: Central strand present. Leaves lanceolate, gradually narrowed to blunt,
acute or acuminate apex, without sheathing base, margins recurved on one or both sides;
costa weakly or rather sharply delimited from leaf lamina, typically with guide cells and
two or rarely only dorsal stereid band, differentiated dorsal and, in several species, also
ventral epidermis; leaf lamina unistratose or with bistratose patches to entirely bistratose
distally; leaf cells rectangular. Rhizoid tubers occasionally present, irregular in shape,
with protruding cells, 100 − 140(−250) × 60 − 95 μm. Dioicous. Perichaetial leaves similar
to lower leaves. Setae red. Capsules inclined, asymmetric, ovoid, gibbous, smooth or fur-
rowed when dry, dark red when mature. Exothecial cells irregular in shape or rectangular,
with thickened longitudinal walls. Annulus weakly differentiated, not revoluble. Peri-
stome dicranoid.
Calcidicranella howei (Renauld & Cardot) Bonfim Santos, Fedosov & Jan Kučera,
comb. nov. ≡ Dicranella howei Renauld & Cardot in Rev. Bryol. 20: 30. 1893.–Type: [United
States of America], Cal. [=California], M.A. Howe.
Calcidicranella varia (Hedw.) Bonfim Santos, Fedosov & Jan Kučera, comb. nov. ≡
Dicranum varium Hedw. in Sp. Musc. Frond.: 133. 1801–Lectotype: [Germany, Leipzig],
[Hedwig?] s.n. (G, Hb. Hedwig-Schwägrichen, barcode G00040364, [36]: Figure 5D–F;
[47]: Figure 77) ≡ Dicranodon varium (Hedw.) Béhéré in Muscol. Rothom. 29. 1826 ≡
Aongstroemia varia (Hedw.) Müll. Hal. in Syn. Musc. Frond. 1: 435. 1848 ≡ Dicranella varia
(Hedw.) Schimp. Coroll. Bryol. Eur. 13: 1856 ≡ Anisothecium varium (Hedw.) Mitt. in J.
Linn. Soc., Bot. 12: 40. 1869.
Calcidicranella pacifica (W.B. Schofield) Jan Kučera & Fedosov, comb. nov. ≡ Di-
cranella pacifica W.B. Schofield, Bryologist 73: 703, 1970.–Holotype: Canada. British Colum-
bia: Vancouver, Spanish Banks, 49°16’N, 123°14′W, seepy silt cliffs and cliff base, Schofield
40,422 (UBC).
Calcidicranella obtusifolia (Berggren) Fedosov, Ignatova & Jan Kučera, comb. et
stat. nov. ≡ Dicranella varia var. obtusifolia Berggren, Kongl. Svenska Vetensk. Acad.
Handl., n.s. 13(7): 36. 1875–Type: Musci Spetsbergens. Exsicc. No. 9. Figure 5 ≡ Anisothe-
cium varium var. obtusifolium (Berggr.) Podp., Consp. Musc. Eur. 118. 1954.

Plants 2023, 12, 1360 21 of 40
Figure 5. Line drawings of gametophyte and sporophyte of Calcidicranella obtusifolia (from: Russia,
Krasnoyarsk Territory, Anabar Plateau, MW9031184, isolate RF42): (A,D) view of fertile plants, dry,
(B) view of fertile plant, wet, (C) capsule, dry, (E) mid-leaf cells, (F) upper-leaf cells, (G,H) leaf trans-
verse sections, (I) exothecial cells, (J–N) stem leaves, (O) perichaetial leaf, (P) basal leaf cells. Scale
bars: 5 mm for (A,B); 2 mm for (C,D); 1 mm for (J–O); 100 μm for (E,F,I,P).
= Anisothecium rubrum var. obtusiusculum Lindb. & Arnell, Kongl. Svenska Vetensk.
Acad. Handl., 23(10): 85. 1890, syn. nov.–Type: ‘Fl. Jen., T. subarct., Polovinka fr.’ [Flora
Jeniseensis, subarctic Taimyr, vicinity of Polovinka River] ≡ Dicranella varia var. obtusi-
uscula (Lindb. & Arnell) Paris, Index Bryol. 336. 1896 ≡ Anisothecium varium var. obtusiuscu-
lum (Lindb. & Arnell) Podp., Consp. Musc. Eur. 118. 1954.
Description: Plants small, gregarious, light green or yellowish. Stems simple, ca. 0.1–
0.2 cm, with strong round central strand and weak sclerodermis, evenly foliated. Leaves
appressed, straight or slightly curved when dry, spreading when moist, 1.0–1.8(–2.2) mm,

Plants 2023, 12, 1360 22 of 40
with wide, ovate bases and more or less distinct shoulders, above shoulders gradually
narrowed towards blunt acumen, concave, lower leaves not widened, triangular; margins
plane at base, narrowly recurved at shoulders and just above and below them or almost
to the leaf tip, unistratose proximally and partly bistratose distally, uneven above, rarely
throughout the margin; costae ending just below apices, rarely percurrent, rather strong,
occupying ca. 1/7–1/5 of leaf base, distinctly delimited from leaf lamina, in transverse sec-
tion with 2–4(–5) large ventral guide cells, differentiated dorsal epidermis and single weak
stereid band; leaf lamina unistratose with occasional bistratose strands distally; upper leaf
cells short rectangular to subquadrate, 12 – 20 × 7 – 12 µm, smooth, not bulging, proxi-
mally longer and wider, 44 – 90 × 10 – 17 µm, elongate-rectangular, 2–3 rows of cells along
margins narrower, ca. 4–6 µm wide. Dioicous. Perichaetial leaves of the same length, but
with wider and longer base, more abruptly narrowed to lanceolate or short subulate acu-
men. Setae 3–5 mm red to brownish. Capsules ca. 1 mm, asymmetric, curved, ovate, with
short neck, strumose, brownish-red, distinctly furrowed, red rimmed distally, exothecial
cells irregular in shape, thick-walled with equally thickened walls, longer and narrower
along furrows, with few stomata proximally. Annulus not differentiated. Operculum
conic. Peristome teeth red to brownish, 450–500 µm long, unequally split for nearly half
of their length, longitudinally striolate proximally, papillose distally. Spores 14–17 µm,
smooth, yellowish-brown, mature in summer. Rhizoidal tubers not seen.
Differentiation: C. obtusifolia resembles C. varia or C. howei in habit but differs in
smaller plants with stems up to 5 mm, while stems of C. varia often extend to 1 cm. Leaf
margins in C. obtusifolia are plane below shoulders, while C. varia has leaf margins re-
curved from the basal leaf portion and C. howei has leaf margins recurved mostly in the
lower leaf part only, often only on one side. Leaf tips in C. obtusifolia are typically blunt,
with costae ending a few cells below tips to being percurrent, while in C. varia/howei leaf
tips are sharp and costae excurrent. Capsules of C. obtusifolia are strumose and distinctly
longitudinally furrowed, while in C. varia/howei capsules are not strumose, smooth or
rarely indistinctly furrowed. Exothecial cells in C. obtusifolia approach C. howei, they are
irregular in shape, with equally thickened walls, while in C. varia longitudinal walls of
exothecial cells typically are thicker than transverse ones. Although in many formal char-
acters C. obtusifolia resembles North American C. pacifica, the latter species is much larger;
moreover, with its contorted to crisped leaves and smooth capsules it is quite distinct from
C. obtusifolia.
Distribution and ecology: A predominantly Arctic species, described from Svalbard
and also known from a single locality in Nenets Autonomous District (European Russia),
suite of localities along Yenisey River, in Taimyr Peninsula, Anabar Plateau and from a
single locality in Yakutia. According to the protologue of Anisothecium rubrum var. obtusi-
usculum [37], it is also one of the most frequent mosses along the Yenisey River banks,
although it rarely occurs in sufficient amounts, while in Svalbard it is either rare or not
recognized from C. varia. It grows on bare loamy soil and silty sediments including saline
ones on eroded slopes along rivers and in massives of baidzarakhs (thermokarst mounds),
most often with Hennediella heimii var. arctica, Funaria spp., Tortula leucostoma, T. cf. cernua,
Bryoerythrophyllum spp., Aloina brevirostris, Stegonia latifolia, Pohlia atropurpurea, Bryum
spp., and many other pioneer mosses. At the same time, according to our field experience,
it differs from other Dicranella s.l. species widespread in Siberian Arctic in occupied habi-
tats, since these usually settle on acidic sandy sediments, typically with gemmiferous spe-
cies of Pohlia, Pogonatum and Psilopilum species.
Protoaongstroemia sachalinensis Fedosov, Ignatova & Jan Kučera, gen. et spec.
nov.–Type: Russian Far East, Sakhalin Island, Tym’ River valley, 50.89518°N, 142.65693°E,
in silty alluvium, 4 September 2009, O.Yu. Pisarenko op03352, MHA (Holotype), MW, NSK
2,003,352 (Isotypes) (Figure 6).

Plants 2023, 12, 1360 23 of 40
Figure 6. Line drawings of gametophyte and sporophyte of Protoaongstroemia sachalinensis (from:
Holotype, isolate FDt122): (A) capsule, dry, (B) perichaetial leaf, (C,H,I) stem leaves, (D) mid-leaf
cells, (E) upper-leaf cells, (F) view of fertile plant, wet, (G) fertile plant, dry, (J) basal leaf cells, (K–
M) leaf transverse sections. Scale bars: 2 mm for (F,G); 1 mm for (B,C,H,I); 0.5 mm for (A); 100 μm
for (D,E,J–M).
Etymology: The generic name originates from Aongstroemia (a genus of dicranoid
mosses) and the prefix proto- (from Greek πρῶτος, first), which reflects the basalmost po-
sition of the genus within the core Aongstroemiaceae clade. The specific epithet reflects
the provenance of the original collection, the Sakhalin Island.
Diagnosis: Differs from other Holarctic Dicranella s.l. species by the combination of
distinctly shouldered leaves, distally regularly 2–3-stratose lamina, costa with a single
stereid band and undifferentiated guide cells, elongate rectangular laminal cells and ir-
regularly furrowed, curved capsules.
Description: Plants small, gregarious, light green or yellowish, mixed with other pi-
oneer mosses. Stems simple, ca. 0.1–0.2 cm, with central strand and weak sclerodermis,
evenly foliated. Leaves appressed, straight or slightly curved when dry, spreading when
moist, gradually increasing in size distally, 1.5 – 1.9 × 0.4 – 0.53 mm, with wide, ovate base,

Plants 2023, 12, 1360 24 of 40
widest at ca. 1/10–1/5 of leaf length with distinct shoulders, abruptly narrowed into grad-
ually tapering blunt acumen, concave; margins plane, with few blunt distant teeth at
shoulders and upper part of acumen to nearly entire, plane, partly bistratose proximally;
costa weak, weakly delimited from leaf lamina, percurrent, in transverse section with ven-
tral and dorsal epidermis and single band of substereids between them, without guide
cells proximally, weakly differentiated distally; leaf lamina unistratose with bistratose
strands proximally, 2–3 stratose distally; leaf cells elongate-rectangular, 37 – 62 × 6 – 13
µm, smooth, bulging on both sides, proximally somewhat longer, 45–75 µm long. Dioi-
cous, male plants not seen. Perichaetial leaves with wider base, abruptly narrowed to
short subulate acumen. Setae reddish, 5–7 mm, spirally twisted when dry and moving
around after wetting. Capsules 1.2–1.5 mm long, asymmetric, curved, ovate, with short
neck, weakly furrowed, not strumose, reddish-brown, red-rimmed distally; exothecial
cells rectangular, moderately thick-walled with evenly incrassate transverse and longitu-
dinal walls, longer and narrower along furrows, with few stomata in proximal part. An-
nulus not differentiated. Operculum conic or with short blunt oblique beak. Peristome
teeth bright red, ca. 300 µm, unequally split for nearly half of their length, longitudinally
striolate proximally, papillose distally. Spores 13–17 µm smooth, yellowish-brown, ma-
ture in autumn. Rhizoidal tubers not seen.
Differentiation: With its shouldered and then gradually narrowed leaves, elongate
rectangular laminal cells and short curved capsules, P. sachalinensis habitually resembles
a small Diobelonella, especially Asian populations with narrower leaves. However, it dif-
fers not only in its size but also in having bistratose leaf lamina. The same trait and plain
margins differentiate P. sachalinensis from the somewhat similar Calcidicranella varia.
Among species with partially bistratose lamina, P. sachalinensis differs from Dicranellopsis
subulata in its non-subulate distal leaf portion, lack of guide cells and undifferentiated
annulus; it differs from Calcidicranella pacifica in having shouldered leaves, narrower and
longer leaf cells and lack of guide cells; and from C. howei in shouldered leaves and nar-
rower costa.
Distribution and ecology: This newly described species is known from a single spec-
imen, which was collected on silty alluvium sediments of Tym’ River in the middle part
of Sakhalin Island. This pioneer moss grew together with Ruficaulis cf. rufescens, Ceratodon
purpureus and male plants of Pohlia cf. lescuriana.
Aongstroemia Bruch & Schimp. in Bryol. Eur. 1: 171 (fasc. 33-36. Mon. 1). 1846, nom.
& orth. cons. ‘Angstroemia’. Type: Aongstroemia longipes (Sommerf.) Bruch & Schimp. [50]
= Anisothecium Mitt., J. Linn. Soc., Bot. 12: 39, 1869, syn. nov.–Type: Anisothecium cam-
pylophyllum (Taylor) Mitt., J. Linn. Soc., Bot. 12: 40. 1869, designated here.
=Dicranella p.pte., Hygrodicranum Cardot p.pte.
? = Eucamptodontopsis Broth. p.pte.
? = Polymerodon Herzog
Note: Since Dicranella now has a conserved type, D. heteromalla (Hedw.) Schimp. [51],
the name Anisothecium should no longer be considered illegitimate. As argued above, the
best candidate to typify the name is Anisothecium campylophyllum with respect to the good
match with the general intent of the author and known phylogenetic affinities of this spe-
cies.
Diagnostic characters: Stem leaves with a broad sheathing base tapering into a blunt
apex (in less developed A. longipes plants) or abruptly narrowed to short or long pointed,
spreading to squarrose leaf apex. Lamina cells rectangular, smooth or sometimes mamil-
lose or papillose, sometimes (irregularly) bistratose. Tubers, if present, spherical without
protruding cells. Capsules erect to inclined, symmetric to asymmetric, oval/obloid to
curved and sometimes slightly strumose, on a straight, erect, red to brownish seta. Annu-
lus not or poorly differentiated.
For a list of accepted species see below.
Excluded species: Aongstroemia filiformis (P. Beauv.) Wijk & Margad. (see above under
Bryopalisotia).

Plants 2023, 12, 1360 25 of 40
Species with uncertain placement: all other accepted species (cf. [22]), pending mor-
pho-molecular studies, and also Aongstroemia orientalis Mitt., for which molecular phylo-
genetic data [10] suggest placement in Ditrichaceae, but additional sampling is required
to assess its affinities within this family.
Aongstroemia boliviana (Herzog) Bonfim Santos & Fedosov, comb. nov. ≡ Hygrodi-
cranum bolivianum Herzog in Biblioth. Bot. 87: 15. pl. 1: f. 1. 1916–Type: [Bolivia] Glazi-
altümpel am Cerro Incachacca, ca. 4600 m, No. 2599; an Steinen im Bach, oberes Llavetal,
ca. 4200 m, No. 4832; in einem Quellbach des Pajonaltales, ca. 4000 m, No. 3264; in einem
Quellbach der Cerros de Malaga, ca. 4000 m, No. 4359.
Aongstroemia campylophylla (Taylor) Müll.Hal. in Syn. Musc. Frond. 2: 608. 1851. ≡
Dicranum campylophyllum Taylor in London J. Bot. 7: 281. 1848–Lectotype (designated in
[39,45]): 8 Aug. 1847 W. Jameson 133 (BM000879353, Isolectotypes BM0006722168,
BM000879354) ≡ Anisothecium campylophyllum (Taylor) Mitt. in J. Linn. Soc., Bot. 12: 40.
1869 ≡ Dicranella campylophylla (Taylor) A. Jaeger in Ber. Thätigk. St. Gallischen Naturwiss.
Ges. 1870-71: 382 (Gen. Sp. Musc. 1: 86). 1872.
=Dicranum cardotii R.Br. bis in Trans. & Proc. New Zealand Inst. 35: 329. 36 f. 9. 1903,
fide [50].–Type: [New Zealand], “on damp banks, tributary of the River Hapuka, near Kai-
koura” Robert Brown s.n. (Holotype: BM-Dixon [52]). ≡ Dicranella cardotii (R.Br. bis) Dixon
in New Zealand Inst. Bull. 3(3): 77. 1923 ≡ Anisothecium cardotii (R. Br. bis) Ochyra in Moss
Fl. King George Island Antarctica 114. 1998.
=Cheilothela vaginata H. Rob. fide [45] = Dicranella convoluta (Hampe) A. Jaeger fide
[45] = Symblepharis tenuis R.S. Williams fide [45].
Aongstroemia canadensis (Mitt.) Siebel & Fedosov, comb. nov. ≡ Cynodontium cana-
dense Mitt., Proc. Linn. Soc., Bot. 8: 17. 1864–Type: [Canada] British N. America (probably
from the Rocky Mountains) T. Drummond, no 101 in part, (probable holotype NY325565)
≡ Dicranella canadensis (Mitt.) Austin in Bot. Gaz. 2: 96. 1877 ≡ Dichodontium canadense
(Mitt.) Lesq. & James in Man. Mosses N. America 62. 1884.
=Dicranella schreberi var. robusta Schimp. ex Braithw. in J. Bot. 9: 289. 1871, syn. nov.–
Type: [United Kingdom, England], at various places in Cheshire, at Milnthorpe (Barnes)
and near Melnrose (c. fr., Jerdon) [specimen Rabenhorst, Bryotheca Europaea No. 74 men-
tioned as bearing well-developed sporophytes ≡ Dicranella schreberiana var. robusta
(Schimp. ex Braithw.) H.A. Crum & L.E. Anderson].
=Anisothecium schreberianum var. elatum (Schimp.) Wijk & Margad. in Taxon 7: 288.
1958, fide [53].
The identity of Dicranella schreberiana var. robusta, treated under this name by [10],
with the type of Cynodontium canadense was suggested by H. Siebel (pers. comm.), who
prepared a detailed account on this taxon.
Aongstroemia grevilleana (Brid.) Müll. Hal. in Syn. Musc. Frond. 1: 439. 1848 ≡ Di-
cranum schreberi var. grevilleanum Brid. in Bryol. Univ. 1: 450. 1826–Type: [UK] In humidis
argillaceis Scotiae. Greville, Hooker, Arnott s.n. ≡ Dicranum grevilleanum (Brid.) Bruch &
Schimp. in Bryol. Eur. 1: 123. 54 (fasc. 37-40. Mon. 19. 7.). 1847 ≡ Dicranella grevilleana (Brid.)
Schimp. in Coroll. Bryol. Eur. 13. 1856 ≡ Anisothecium grevilleanum (Brid.) Arnell & C.E.O.
Jensen in Bih. Kongl. Svenska Vetensk.-Akad. Handl. 21 Afd. 3(10): 49. 1896 ≡ Dicranella
schreberi var. grevilleana (Brid.) Mönk. in Laubm. Eur. 179. 1927.
Aongstroemia herrerae (R.S. Williams) Bonfim Santos & Fedosov, comb. nov. ≡ Hy-
grodicranum herrerae R.S. Williams in Bryologist 29: 37. pl. 3: f. 1–9. 1926 (‘herrerai’, cf. ICN
Art. 60.8.a)–Type: “Growing about waterfalls, Río Tapfi, province of Cuzco, Peru, at 3600
m.” F.L. Herrera No. 792, Sept. 1925, same locality, F.L. Herrera No. 798a”.
Aongstroemia hookeri Müll.Hal., Syn. Musc. Frond. 2: 607. 1851.–Type: Insula Er-
emitae ad Cap. Horn: J.D. Hooker. ≡ Anisothecium hookeri (Müll. Hal.) Broth., Nat. Pflan-
zenfam. (ed. 2) 10: 178. 1924. ≡ Dicranella hookeri (Müll. Hal.) Cardot, Bull. Herb. Boissier,
sér. 2, 6: 4. 1906.
=Anisothecium perpusillum Dusén fide [54] = Dicranella subclathrata Lorentz fide [55] =
Meesia patagonica Dusén fide [55].

Plants 2023, 12, 1360 26 of 40
? = Polymerodon andinus Herzog, Beih. Bot. Centralbl., 26(2): 48. pl. 1. 1909.–Type: Bo-
livia: An feuchten Felsen neben dem Weg im Valle de Llave (bei Cochabamba), ca. 3600
m, mit Wollnya stellata Herzog: Januar, 08.
? = Eucamptodontopsis pilifera (Mitt.) Broth., Nat. Pflanzenfam. (ed. 2) 10: 202. 1924. ≡
Eucamptodon pilifer ‘piliferus’ Mitt., J. Linn. Soc., Bot., 12: 69. 1869.–Type: Trinidad, Marga-
rita, Palma Real, Crüger.
Aongstroemia longipes (Sommerf.) Bruch & Schimp., Bryol. Eur. 1: 173 (fasc. 33–36
Monogr. 3). 1846. ≡ Weissia longipes Sommerf., Suppl. Fl. Lapp. 52, pl. 1, f. 1–10. 1826.–
Type: “In terra argillosa humida ad rivulos montanos in provincia Saltdalen Norvegiae
(Sommerfelt s.n.); in Canada superiore (Drummond s.n.)”.
Aongstroemia schreberiana (Hedw.) Bonfim Santos & Fedosov, comb. nov. ≡ Di-
cranum schreberianum Hedw. in Sp. Musc. Frond. 144, pl. 33, f. 6-10. 1801–Lectotype: [Ger-
many, Saxony, Leipzig], sin. coll. s.n. (G, Hb. Hedwig-Schwägrichen, barcode G00040018,
[36]: Figure 3F–H; [47]: Figure 73) ≡ Anisothecium schreberianum (Hedw.) Dixon in Rev.
Bryol. Lichénol. 6: 104. 1934 ≡ Dicranella schreberiana (Hedw.) Hilf. ex H.A. Crum & L.E.
Anderson in Mosses E. N. Amer. 1: 169. 1981 = Bryum crispum Schreber, nom. inval., Spic.
Fl. Lips. 79 (no. 1038). 1771 = Dicranum schreberi Sw. nom. illeg., Monthly Rev. 34: 538. 1801.
5. Materials and Methods
5.1. Taxon Sampling
The matrix of molecular data was largely based on that used for the backbone phy-
logeny of Dicranidae, with a focus on Dicranella and Aongstroemia [10]. With respect to the
absence of dicranelloid taxa in some lineages of haplolepidous mosses, we reduced the
matrix by leaving out or reducing the number of accessions in lineages where these taxa
were absent, in order to decrease the complexity of the alignment. The outgroups were
thus reduced to include only Pseudoditrichum, Flexitrichum, Scouleria, Drummondia, Hyme-
noloma, and Bryoxiphium, and we further substantially reduced the representation of Leu-
cobryaceae, Rhabdoweisiaceae, Dicranaceae, and related families (leaving out completely
Mittenia, Pleurophascum, Serpotortella, Hypodontiaceae, Octoblepharaceae, and Calym-
peraceae), and also Ditrichaceae including Aongstroemia orientalis and A. julacea, which
will be treated in a dedicated future article. On the other hand, we added accessions of
Dicranella staphylina, D. humilis, D. varia incl. its neglected var. obtusifolia, D. pacifica, D.
grevilleana, D. schreberiana incl. its var. robusta, D. campylophylla, D. hookeri, D. heteromalla,
D. curvipes, D. polii, D. vaginata, Aongstroemia longipes, A. filiformis, Neodicranella hamulosa,
and unassigned dicranelloid plants from Pacific Russia, which were found to be related
to Dicranella s.str. and to Diobelonella/Dichodontium/Neodicranella grade. Newly generated
trnK–psbA data were added for at least one representative of each major lineage left. La-
boratory protocols for isolation of DNA, amplification and sequencing followed the pro-
tocols described in [8,10,56,57]. Genbank accession numbers of the included specimens
and vouchers of specimens studied de novo are compiled in Appendix A.
5.2. Phylogenetic Analyses
Sequences were aligned using MAFFT v. 7 [58] with the E-INS-i strategy and other-
wise default settings, and the resulting alignment was improved manually at obviously
misaligned sites. The concatenated dataset (available in http://purl.org/phylo/tree-
base/phylows/study/TB2:S30163) was tentatively partitioned according to the sequenced
regions (trnF–trnL, abbreviated L hereafter, trnL–rps4 (T), rps4–trnS (R), trnK–psbA (K),
nad5 (N) with respect to their significantly differing coverage, rather than according to
coding and non-coding regions. The best-fit partitioning scheme and models of nucleotide
evolution were searched for in PartitionFinder2 [59]. The results of the greedy algorithm
used suggested partitioning according to all of the initially suggested partitions, with the
HKY + I+G model for the trnF–trnL partition and GTR + I+G for the remaining ones. Indel
data were scored for individual partitions using the simple indel coding (SIC) approach

Plants 2023, 12, 1360 27 of 40
[60] in SeqState 1.4.1 [61] and added to the dataset in three variants: (1) indels scored only
for L, R, and N partitions; (2) indels scored for L, T, R, and N partitions; and (3) indels
scored for all partitions. Based on the results of [10], we did not separately analyze the L,
R, and N data with respect to the reported absence of conflicts in topology and relatively
low resolution of trees obtained from single-gene analyses, but we explored the influence
of previously unused regions, i.e., (a) data from spacers flanking trnT between trnL and
rps4 (T) and (b) trnK–psbA (K) data, which were successively added to the working pilot
analyses. Given the amount of phylogenetic signal, the K data were also analyzed sepa-
rately from the 52 accessions for which these data were available.
Phylogenetic reconstructions were performed using Bayesian inference (BI) and max-
imum likelihood (ML). BI was run in MrBayes v.3.2.7 [62] in two parallel runs, each con-
sisting of eight Markov chains run for 2,000,000 generations as default, and with further
generations added if the convergence between runs did not reach 0.01, with the default
number of swaps and a sampling frequency of one tree for each 100 generations. The chain
temperature was initially set at 0.1 and lowered as necessary according to the acceptance
rates. The models were sampled throughout the GTR model space and gamma-distrib-
uted rate variation across sites, and a proportion of invariable sites, as suggested by the
PartitionFinder. PSRF values, were checked as being close to 1.000. ESS values were
checked using Tracer v.1.7.2 [63] as being higher than 200. Consensus trees were calcu-
lated after omitting the burn-in of the first 25% of trees. The best-scoring maximum like-
lihood (ML) trees were searched using the new rapid hill-climbing algorithm in RAxML
8.2.12 [64] under the GTR model with gamma model of rate heterogeneity in 50 independ-
ent runs, each starting from a different random tree. The extended majority-rule consen-
sus tree criterion was used to stop the bootstrapping used for the assessment of the node
robustness. Analyses were performed using the grid computational services provided by
the MetaCentrum Virtual Organization (see Acknowledgement). Trees were visualized
using TreeGraph2 [65].
5.3. Morphological Studies
In addition to standard microscopic observations during the revision of herbarium
specimens, preparation of taxon descriptions, and illustrations, images of peristomes were
obtained by scanning electron microscopy (SEM) with a JSM-6380 (JEOL) at the User Fa-
cilities Center of M.V. Lomonosov Moscow State University. Peristomes mounted on
stubs were coated with gold without any additional preparation, and light microscope
illustrations were made under a stereomicroscope Olympus SZX-7 with a digital camera
Infinity 8, with Z-stacking in Helicon Software [66].
Supplementary Materials: The following supporting information can be downloaded at:
https://www.mdpi.com/article/10.3390/plants12061360/s1. Figure S1: Maximum Likelihood tree, in-
ferred from the concatenated data matrix from the chloroplast trnF–trnS and mitochondrial nad5
intron 2 sequence alignment (LTRN dataset) of selected species of Dicranidae. Bootstrap support
values inferred from ML analyses without and with indel coding are shown above branches, poste-
rior probabilities inferred from BI without and with indel coding are shown below branches. The
same tree with saved branch lengths is shown in the lower left corner. For details, see File S1; Figure
S2: Maximum Likelihood tree, inferred from and trnK–psbA of selected species of Dicranidae. Boot-
strap support values inferred from ML analyses without and with indel coding are shown above
branches, posterior probabilities inferred from BI without and with indel coding are shown below
branches. The same tree with saved branch lengths is shown in the lower left corner. For details, see
File S1.
Author Contributions: Conceptualization, V.F. and J.K.; Methodology, V.F., A.F., and J.K.; Valida-
tion, E.I. and J.K.; Formal analysis, J.K.; Investigation, V.F., A.F., E.I., and J.K.; Resources, V.F. and
J.K.; Data curation, E.I. and J.K.; Writing—original draft, V.F., E.I., and J.K.; Visualization, V.F. and
E.I. All authors have read and agreed to the published version of the manuscript.

Plants 2023, 12, 1360 28 of 40
Funding: The work of V.E. Fedosov, E.A. Ignatova, and A.V. Fedorova was supported by Grant #
14-18-00121 and 14-18-00121П from the Russian Science Foundation (RNF). We also thank the Min-
istry of Higher Education and Science of the Russian Federation for support and the Center of Col-
lective Use “Herbarium MBG RAS” (grant 075-15-2021-678). Molecular analyses performed by Jan
Kučera at the University of South Bohemia were funded by the institutional sources of the Faculty
of Science. Computational resources (‘Metacentrum VO’) were supplied by the Ministry of Educa-
tion, Youth and Sports of the Czech Republic under the Projects CESNET (Project No. LM2015042).
The work of A.V. Fedorova was also supported by Tsitsin Main Botanical Garden state assignment
no. 122042500074-5.
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: All authors agree with MDPI Research Data Policies.
Acknowledgments: First of all, we are grateful to Marina Bonfim Santos and Michael Stech (Natu-
ralis Biodiversity Center and Leiden University, Leiden, The Netherlands) as well as Henk Siebel
(Natuurmonumenten, Hilversum, The Netherlands) for permission to use the results of their study.
We acknowledge the molecular laboratory work performed by Dr. Alžběta Manukjanová (Univer-
sity of South Bohemia) and curators of herbaria, which provided specimens for DNA extraction and
the morphological study (B, CONC, DUKE, LE, MHA, NSK, MO, NY). The work on SEM was per-
formed under the financial support of the Ministry of Education and Science of the Russian Feder-
ation. We also are grateful to Juan Larraín for useful comments on the identity of several south-
Hemispheric taxa and to John Brinda for checking the syntype specimen of Dicranella varia var. ob-
tusifolia in MO.
Conflicts of Interest: The authors declare no conflict of interest.

Plants 2023, 12, 1360 29 of 40
Appendix A
Studied specimens and GenBank accession numbers. Specimens are annotated with the isolate codes shown in the trees (indicated as “N/A”
for sequences for which “isolate” annotation is omitted in GenBank), geographic origin and specimen ID (where available). Isolate codes of the
originally studied specimens are in bold. Each isolate starts with a new line, so different isolates of the same species combined in the concatenated
datasets are indicated with an “&” mark after the isolate code, which means that the present sequence(s) was combined with the one on the next
line.
Isolate
Organism
Country
Specimen Voucher
trnL-trnF
rps4-trnL
trnS-rps4
trnK
Nad5
PSD
Pseudoditrichum mirabile
Russia: Taimyr Eriechka
River upper course
Fedosov 13-3-1028 MW
OQ199090
OQ199090
OQ199090
OQ199854
KR026964
Fx2975
Flexitrichum flexicaule
Czech Republic:
Vápenná
Kučera 19465 (CBFS)
OQ094918
OQ094918
OQ094918
OQ094950
OQ094881
Fx2978
Flexitrichum gracile
Czech Republic: Horní
Albeřice
Kučera 22345 (CBFS)
OQ094919
OQ094919
OQ094919
-
-
N/A &
Drummondia prorepens
Allen 6192
JQ690728
JQ690728
-
-
-
MS1011 &
Drummondia prorepens
-
-
KX580512
-
-
B137 &
Drummondia prorepens
-
-
-
LN828317
-
N/A
Drummondia prorepens
Quandt B137b
-
-
-
-
EU095282
OK757
Scouleria pulcherrima
Russia: Irkutsk Province
Mamontov LE321
KX446936
-
KX369282
OQ199855
KX369284
Hl1219
Hymenoloma crispulum
Austria: Tyrol, Mt Hoher
Riffler
Kučera 18871 (CBFS)
OQ094921
OQ094921
OQ094921
OQ094952
OQ094883
Bx2964
Bryoxiphium japonicum
Russia: Primorsky Terri-
tory, Mt Pidan
Kučera 21746 (CBFS)
OQ094895
OQ094895
OQ094895
OQ094934
OQ094864
Dl2913
Dicranella staphylina
Czech Republic: Kaplice,
Střítež, Malý Chuchelec
Kučera 22785 (CBFS)
OQ094911
OQ094911
OQ094911
OQ094943
OQ094874
TJH05
Dicranella staphylina
Netherlands
Aptroot 69818 (L)
MN178095
-
MN187514
-
MN178020
TJH27
Dicranella staphylina
Netherlands
Siebel 2013.451 (L)
MN178096
-
MN187515
-
MN178021
FDt107
Dicranella staphylina
Russia: Krasnoyarsk Ter-
ritory, Putorana Plateau
MW9117945
OQ199091
-
OQ199091
OQ199856
OQ199058
FDt116
Dicranella staphylina
Russia: Krasnoyarsk Ter-
ritory, Putorana Plateau
MW9117878
OQ199092
-
OQ199092
-
-
BF59
Dicranella staphylina
Russia: Yakutia
MHA9049571
OQ199093
-
OQ199093
-
OQ199059

Plants 2023, 12, 1360 30 of 40
BF63
Dicranella staphylina
Russia: Khabarovsk Ter-
ritory, Badzhal
Pisarenko, NSK (as D. hu-
milis), dupl. MW
OQ199094
-
OQ199094
-
OQ199060
BF64
Dicranella staphylina
Russia: Khabarovsk Ter-
ritory, Badzhal
Fedosov, MW9130183
OQ199095
-
OQ199095
-
OQ199061
Ar2959
Archidium alternifolium
Czech Republic: Zvůle
Kučera 23203 (CBFS)
OQ094892
OQ094892
OQ094892
OQ094931
OQ094861
Lm2476 &
Leucobryum glaucum
Czech Republic: Žofinka
reserve
Kučera 23239 (CBFS)
OP081669
-
-
-
-
B966 &
Leucobryum glaucum
M. Krug, B966
-
-
-
LN828327
-
N/A
Leucobryum glaucum
Muhle 281097-6 (ULM)
-
-
-
-
AJ291560
MDP336 &
Brothera leana
Long DG 21998 April 19 1992
(DUKE)
-
-
AY908129
-
AY908911
N/A &
Brothera leana
HIRO120176
-
-
-
AB742510
-
CB52
Brothera leana
China
FJ572426
-
-
-
-
Sa2977
Saelania glaucescens
Czech Republic: Krko-
noše Mts. Čertova
zahrádka
Kučera 23757 (CBFS)
OQ094925
OQ094925
OQ094925
OQ094955
OQ094886
BG577 &
Ptychomitrium gardneri
Ireland 7038 (PMAE)
-
-
AY908616
-
AY908951
N/A &
Ptychomitrium gardneri
AF023719
-
-
-
-
N/A &
Ptychomitrium gardneri
-
EU246894
-
-
-
MO5135689
Ptychomitrium gardneri
MO5135689
-
-
-
KX024249
-
Bd2962
Brachydontium trichodes
Czech Republic: Krko-
noše Mts. Čertova louka
Kučera 23844 (CBFS)
OQ094894
OQ094894
OQ094894
OQ094933
OQ094863
Gr3041
Grimmia pulvinata
Czech Republic:
Rašovické skály
Kučera 20140 (CBFS)
OQ094920
OQ094920
OQ094920
OQ094951
OQ094882
N/A &
Racomitrium lanuginosum
KRAM B 43266
MN239148
-
MN239160
-
-
N/A &
Racomitrium lanuginosum
Jorgensen et al. 1589 (MA)
-
HE586592
-
HE588126
-
N/A
Racomitrium lanuginosum
Muhle 070997-10 (ULM)
-
-
-
-
EU095285
Bl2960
Blindia acuta
Czech Republic: Krko-
noše Mts. Čertova
zahrádka
Kučera 23747 (CBFS)
OQ094893
OQ094893
OQ094893
OQ094932
OQ094862
RF55
Dicranella subulata
Russia: East Siberian
North
MW9007554
MN178097
-
MN187516
-
MN178022
TJH08
Dicranella subulata
Italy
Siebel 2015.313 (L)
MN178098
-
MN187517
-
MN178023

Plants 2023, 12, 1360 31 of 40
TJH19
Dicranella subulata
Norway
Siebel 2014.610 (L)
MN178099
-
MN187518
-
MN178024
TJH34
Dicranella subulata
Italy
Siebel 2015.357 (L)
MN178100
-
MN187519
-
MN178025
Dl2969
Dicranella subulata
Czech Republic: Krko-
noše Mts., Rudný potok
Kučera 23770 (CBFS)
OQ094912
OQ094912
OQ094912
OQ094944
OQ094875
RF45
Dicranella crispa
Russia: East Siberian
North
MW9007542
MN178069
-
MN187488
-
MN177999
TJH01
Dicranella crispa
Norway
Siebel 2014.765 (L)
MN178070
-
MN187489
-
MN178000
Dl2966
Dicranella crispa
Sweden: Lule Lappmark,
Kamajokk
Kučera 19719 (CBFS)
OQ094897
OQ094897
OQ094897
OQ094936
OQ094866
FDt115
Dicranella humilis
Czech Republic:
Rýmařov
Koval s.n. (CBFS:14897)
OQ199096
-
OQ199096
-
-
BF71
Dicranella humilis
Russia: Leningrad Prov
LE
OQ199097
-
OQ199097
-
OQ199062
FDt119
Dicranella cf. rufescens
Russia: Sakhalin
op03352 (MHA)
OQ199098
-
OQ199098
-
OQ199063
RF63
Dicranella cf. rufescens
Russia: Ural Mts.
MW9030966
MN178088
-
MN187507
-
MN178014
RF62
Dicranella rufescens
Russia: West Siberia
MW9030986
MN178087
-
MN187506
-
MN178013
BCNL2
Dicranella rufescens
Netherlands
Smulders 08151 (L0872909)
MN178086
-
MN187505
-
MN178012
BCNL1
Dicranella rufescens
Netherlands
Pellicaan s.n. (L0873203)
MN178085
-
MN187504
-
-
Dl2967
Dicranella rufescens
Czech Republic: Šumava
Mts., Nová Pec, Říjiště
Kučera 23913 (CBFS)
OQ094908
OQ094908
OQ094908
OQ094941
OQ094872
RF84
Amphidium lapponicum
Russia: Kola Peninsula
MW9078004
MN092374
-
MN092437
OQ199857
MN092571
49156
Amphidium sp.
49156
MW429503
MW429503
MW429503
MW429503
-
Sc2980
Schistostega pennata
Czech Republic: Krko-
noše Mts. Čistá valley
Kučera 21209 (CBFS)
OQ094926
OQ094926
OQ094926
OQ094956
OQ094887
Dw2974
Dicranoweisia cirrata
Czech Republic:
Řevničov
Kučera 23389 (CBFS)
OQ094914
OQ094914
OQ094914
OQ094946
OQ094877
Ki1562
Kiaeria starkei
Czech Republic: Krko-
noše Mts. Studniční hora
Kučera 19081 (CBFS)
OQ094922
OQ094922
OQ094922
OQ094953
OQ094884
Tn2981
Trematodon ambiguus
Norway: Nordland,
Rana, Svartisvatnet
Kučera 6965 (CBFS)
OQ094927
OQ094927
OQ094927
OQ094957
OQ094888
Rf2976
Rhamphidium purpuratum
Greece: Crete, Chania
Blockeel 40/147 (CBFS)
OQ094924
OQ094924
-
-
-
MBS135
Rhamphidium purpuratum
Portugal, Azores
Stech 08-392 (L)
MN178117
-
MN187538
-
MN178044
MDP238
Rhamphidium dicranoides
Ramirez B.R. et al. 8.339A Aug
23 1995 (MO)
-
-
AY908089
-
AY908867

Plants 2023, 12, 1360 32 of 40
N/A
Symblepharis krausei
Larrain 42471 (CONC)
MN718488
-
MN718549
-
MN718519
BF74
Dicranella vaginata
Chile
Ignatov & Ignatova 2019 (MW)
OQ199099
-
OQ199099
-
OQ199064
FDt194
Dicranella vaginata
Chile
Larraín 41567 (MW)
OQ199100
OQ199100
OQ199100
-
OQ199065
FDt195
Dicranella vaginata
Chile
Larraín 39297 (MW)
OQ199101
OQ199101
OQ199101
-
OQ199066
Td2932
Trichodon cylindricus
Czech Republic: Krko-
noše Mts., Mt. Kotel
Kučera 22223 (CBFS)
OQ094928
OQ094928
OQ094928
-
OQ094889
B894
Trichodon cylindricus
M. Krug
-
-
-
LN828325
-
BR_13622234 &
Ceratodon purpureus
France
BR 5040136222346 Bamps
P.4685
MN552345
-
-
-
-
MSCp &
Ceratodon purpureus
-
-
KX580500
-
AY908862
B893
Ceratodon purpureus
-
-
-
LN828324
-
N/A
Ditrichum rhynchostegium
Y. Inoue 5462 (HIRO)
LC716918
LC716918
LC716918
LC716918
LC716919
N/A
Scopelophila cataractae
Y. Inoue 4216 (HIRO)
LC634773
LC634773
LC634773
LC634773
LC634774
N/A
Pseudocrossidium replicatum
coll. Miguel Angel Villalobos
NC_056241
NC_056241
NC_056241
NC_05624
1
MT310681
G2480
Gymnostomum calcareum
Greece: Central Macedo-
nia, Litochoro, Enipeas
valley
Kučera 19624 (CBFS)
OQ199102
OQ199102
OQ199102
OQ199858
OQ199067
KBDI00015
Fissidens nobilis
NC_044155
NC_044155
NC_044155
NC_04415
5
-
Fi3037
Fissidens bryoides
Czech Republic: Bítov
Kučera 24319 (CBFS)
OQ094917
OQ094917
OQ094917
OQ094949
OQ094880
N/A
Chorisodontium aciphyllum
MW355440
MW355440
MW355440
MW355440
NC_050308
Di3039
Dicranum scoparium
Czech Republic: Krko-
noše Mts. Kotelský potok
Kučera 21176 (CBFS)
OQ094915
OQ094915
OQ094915
OQ094947
OQ094878
FDt148
Chrysoblastella chilensis
Chile: Valdivia
MW9113147
OQ199103
-
OQ199103
OQ199859
-
MSCc
Chrysoblastella chilensis
AF135097
-
KX580501
-
KX580439
MBS148
Bryowijkia madagassa
Madagascar
Magill & al. 9975 (L)
MN178055
-
MN187476
-
-
B14
Bryowijkia ambigua
Ellis 901 (BM)
-
-
AY908100
-
AY908873
MDP492
Eccremidium floridanum
Allen 7505 (DUKE)
-
-
AY908098
-
AY908872
MDP68
Cladophascum gymnomitrioides
Perold 2475 (MO)
-
-
AY908097
-
AY908871
Ao2890
Aongstroemia filiformis
Madagascar: Vakona Re-
serve
Brinda 12381 (CBFS)
OQ094890
OQ094890
OQ094890
OQ094929
OQ094859

Plants 2023, 12, 1360 33 of 40
MBS157
Aongstroemia filiformis
Reunion
Schäfer-Verwimp & Verwimp
s.n. (MA)
MN178046
-
-
-
-
N/A
Aongstroemia filiformis
Allen 6403 (DUKE)
-
-
AY908094
-
AY908869
RF74
Aongstroemia filiformis
Colombia
MA-Musci-29275 (duplicate in
LE)
MN178047
-
MN187469
-
-
FDt39
Microcampylopus sp.
Australia: New South
Wales
Streimann 63666 (as Ditrichum
difficile) (LE)
OQ199104
-
OQ199104
-
OQ199068
MSMl2
Microcampylopus laevigatus
Reunion
Greven & Khoeblal 4000/12 (L)
MN178115
-
MN187535
-
MN178041
MSMf
Leptotrichella flaccidula
Schultze-Motel 3209 (B)
-
-
KX580520
-
KX580400
Dl2915
Dicranella polii
Madagascar: Vakona Re-
serve
Brinda 12371 (CBFS)
OQ094907
OQ094907
OQ094907
OQ094940
OQ094871
FDt33
Garckea flexuosa
Australia: Melville Island
Streimann 42410 (LE)
-
-
OQ199105
-
-
MDP124
Garckea phascoides
Magill & Pocs 11583 (MO)
-
-
AY908096
-
AY908870
MS1031
Microcampylopus sp.
Salvador
Bacaro s.n. (ITIC, dupl. in L)
MN178114
-
MN187533
-
MN178039
MBS48
Campylopodium medium
Lesser Sunda Islands
Schmutz 6706 (L)
MN178056
-
MN187477
-
-
MSMk
Microcampylopus khasianus
-
-
KX580527
-
KX580412
Mc2036
Microcampylopus khasianus
Cameroon: Mt Cameroon
Dančák et al. (CBFS22732)
OQ094923
OQ094923
OQ094923
OQ094954
OQ094885
MSMcu
Microcampylopus curvisetus
Venezuela
Schäfer-Verwimp & Verwimp
12351 (L)
-
-
MN187534
-
MN178040
FDt5
Dicranella cerviculata
Russia: Kamchatka Pen-
insula
MW9030767
MN178063
-
MN187483
-
MN177995
FDt6
Dicranella cerviculata
Russia: North Siberia
MW9036970
MN178064
-
MN187484
-
MN177996
TJH07
Dicranella cerviculata
Netherlands
Aptroot 69861 (L)
MN178066
-
MN187486
-
-
RF61
Dicranella cerviculata
Russia: Far East
MW 9030770
MN178065
-
MN187485
-
MN177997
RF49
Dicranella heteromalla
Russia: Vladimir Prov-
ince
MW9030717
OQ199106
-
OQ199106
-
OQ199069
RF47
Dicranella heteromalla
Russia: Vladimir Prov-
ince
MW9030879
MN178079
-
MN187498
-
MN178007
FDt35
Dicranella heteromalla
Russia: Primorsky Terri-
tory
MHA9109527 (as Ditrichum
macrorrhynchum)
OQ199107
-
OQ199107
-
OQ199070
Dl2536
Dicranella heteromalla
Russia: Primorsky Terri-
tory, Chandolaz
Kučera 21639 (CBFS)
-
OQ094903
OQ094903
-
-
BG1366
Dicranella heteromalla
Goffinet 8162 (CONN)
-
-
AY908099
-
AY908938

Plants 2023, 12, 1360 34 of 40
FDT128
Dicranella heteromalla
Chile
Larraín 39274 (MW)
OQ199108
-
OQ199108
-
-
TJH28
Dicranella heteromalla
Netherlands
Buter s.n. (L0873082)
MN178081
-
MN187500
-
MN178009
TJH12
Dicranella heteromalla
Netherlands
Zwarts s.n. (L0873198)
MN178080
-
MN187499
-
MN178008
SF109
Dicranella heteromalla
Chile: Falklands
Larraín 39210 (MW)
OQ199109
-
OQ199109
-
-
RF46
Dicranella heteromalla
Norway
MW 9030851
MN178078
-
MN187497
-
-
Dl2601
Dicranella heteromalla
Czech Republic: Krko-
noše Mts. Hrazený důl
Kučera 23556 (CBFS)
OQ094904
OQ094904
OQ094904
OQ094938
OQ094869
Dl2995
Dicranella curvipes
Russia: Irkutskaya Prov-
ince, Snezhnaya River
Kučera 20616 (CBFS)
OQ094898
OQ094898
OQ094898
-
-
Dl3047
Dicranella curvipes
Russia: Primorsky Terri-
tory, Mt Pidan
Kučera 21778 (CBFS)
OQ094899
OQ094899
OQ094899
-
-
Dl3049
Dicranella curvipes
Russia: Primorsky Terri-
tory, Elomovsky Klyuch
Kučera 21379 (CBFS)
OQ094900
OQ094900
OQ094900
-
-
RF58
Dicranella cf. curvipes
Russia: Far East
MW9030958
MN178071
-
MN187490
-
MN178001
RF59
Dicranella cf. curvipes
Russia: Far East
MW9030952
MN178072
-
MN187491
-
MN178002
RF60
Dicranella curvipes
Russia: Far East
MW9030946
MN178073
-
MN187492
-
-
CF1_1
Dicranella thermalis
Russia: Iturup Island
MW9090383
OQ199110
-
OQ199110
-
OQ199071
CF1_2
Dicranella thermalis
Russia: Iturup Island
MW9073557
OQ199111
-
OQ199111
-
OQ199072
Dl3061
Dicranella pacifica
Canada: British Colum-
bia, Kaien Island
Schofield & Sharp 25837 (CAS)
-
OQ094905
OQ094905
-
-
Dl3062
Dicranella pacifica
Canada: British Colum-
bia, Moresby Island
Schofield & Sharp 25944 (CAS)
OQ094906
OQ094906
OQ094906
OQ094939
OQ094870
TJH02
Dicranella howei
Netherlands
Siebel 2014.155
MN178082
-
MN187501
-
MN178010
TJH06
Dicranella howei
France
Bijlsma 12266 (L)
MN178083
-
MN187502
-
-
TJH30
Dicranella howei
Greece
Nieuwkoop 2015559
(L0255415)
MN178084
-
MN187503
-
MN178011
BF45
Dicranella varia
Russia: Ingushetia
MW9090785
OQ199112
-
OQ199112
-
OQ199073
RF56
Dicranella varia
Belarus
MW9031125
MN178102
-
MN187521
-
MN178027
TJH29
Dicranella varia
Netherlands
Siebel 2015.531 (L)
MN178103
-
MN187522
-
MN178028
TJH36
Dicranella varia
Belgium
Siebel 2015.440 (L)
MN178104
-
MN187523
-
MN178029
BF48
Dicranella varia
Finland
MW9078071
OQ199113
-
OQ199113
-
OQ199074
BF56
Dicranella varia
Russia: Pskov Province
MHA9049664
OQ199114
-
OQ199114
-
OQ199075
BF57
Dicranella varia
Russia: Altai
MHA9049545
OQ199115
-
OQ199115
-
OQ199076

Plants 2023, 12, 1360 35 of 40
BF60
Dicranella varia
Russia: Sakhalin Island
MHA9049544
OQ199116
-
OQ199116
-
OQ199077
BF53
Dicranella varia
Russia: Krasnoyarsk
Terr. Putorana Plateau
MW9007559
OQ199117
-
OQ199117
-
OQ199078
Dl2971
Dicranella varia
Czech Republic: Dolní
Červená Voda
Kučera 23946 (CBFS)
OQ094913
OQ094913
OQ094913
OQ094945
OQ094876
RF42
Dicranella obtusifolia
Russia: Krasnoyarsk
Terr. Anabar Plateau
MW9031184
MN178101
-
MN187520
-
MN178026
BF54
Dicranella obtusifolia
Russia: Krasnoyarsk
Terr. Taimyr, Ary-Mas
MW9031190
OQ199118
-
OQ199118
-
OQ199079
BF58
Dicranella obtusifolia
Russia: Yakutia
MHA9049564
OQ199119
-
OQ199119
-
OQ199080
BF46
Dicranella obtusifolia
Russia: Krasnoyarsk
Terr. Taimyr, Dickson
MW9114171
OQ199120
-
OQ199120
-
OQ199081
FDt122
Protoaongstroemia sacha-
linensis
Russia: Sakhalin Island
op03352 MHA (admixture)
OQ199121
-
OQ199121
OQ199860
OQ199082
DF10
Diobelonella palustris
Russia: Iturup Island Pa-
rusnaya Bay
MW9007534
OQ199122
-
OQ199122
-
OQ199083
Db2945
Diobelonella palustris
Czech Republic: Krko-
noše Mts. Velka Kotelní
jáma
Kučera 22284 (CBFS)
OQ094916
OQ094916
OQ094916
OQ094948
OQ094879
Dc2941
Dichodontium pellucidum
Czech Republic: Jablonec
nad Jizerou, Hradsko
Kučera 15658 (CBFS)
OQ094896
OQ094896
OQ094896
OQ094935
OQ094865
Dc2973
Dichodontium flavescens
United Kingdom: Scot-
land, vc89, Pitlochry
Kučera 10090 (CBFS)
-
OQ199123
OQ199123
-
-
BF70
Dichodontium cf. flavescens
Russia: Shikotan Island
Fedosov 2021 MW
OQ199124
-
OQ199124
-
OQ199084
IPG1
Dichodontium flavescens
Luxembourg
Siebel 2012.223 (L)
MN178059
-
MN187479
-
-
BF9
Neodicranella hamulosa
Portugal
Porley 17.V.2020 (dupla in
MW)
MW798729
-
MW798725
OQ199861
MW798721
BF10
Neodicranella hamulosa
Portugal
Porley 20.I.2019 (dupla in
MW)
-
-
MW798726
-
MW798722
BF11
Neodicranella hamulosa
Portugal
Porley 30.I.2017 (dupla in
MW)
MW798730
-
MW798727
-
MW798723
Ao2924
Aongstroemia longipes
Austria: Carinthia,
Sandersee
Kučera 12803 (CBFS)
OQ094891
OQ094891
OQ094891
OQ094930
OQ094860

Plants 2023, 12, 1360 36 of 40
MBS154
Aongstroemia longipes
Norway
Brand s.n. (herb. H.J. During)
MN178048
-
MN187470
-
MN177982
RF43
Aongstroemia longipes
Russia: North Siberia
MW9002156
MN178050
-
MN187471
-
MN177984
Dl2912
Dicranella grevilleana
Norway: Troms og Finn-
mark, Målselv, Návsti
Kučera 15895 (CBFS)
OQ094901
OQ094901
OQ094901
OQ094937
OQ094867
Dl2916
Dicranella grevilleana
Austria: Carinthia,
Sandersee
Kučera 12802 (CBFS)
OQ094902
OQ094902
OQ094902
-
OQ094868
RF54
Dicranella grevilleana
Russia: Yakutia
MW9074884
MN178076
-
MN187495
-
MN178005
RF38
Dicranella grevilleana
Russia: Far East, Sakhalin
Island
MW9030841
MN178074
-
MN187493
-
MN178003
RF39
Dicranella grevilleana
Russia: North Siberia
MW9030839
MN178075
-
MN187494
-
MN178004
TJH25MBS150
Dicranella grevilleana
Norway
Siebel 2012.291 (L)
MN178077
-
MN187496
-
MN178006
Dl2923
Dicranella schreberiana
Czech Republic: Horní
Albeřice
Kučera 22365 (CBFS)
OQ094909
OQ094909
OQ094909
OQ094942
OQ094873
Dl2926
Dicranella schreberiana
Greece: Epirus, Ioannina,
Pades
Kučera 19559 (CBFS)
OQ094910
OQ094910
OQ094910
-
-
RF40
Dicranella cf. schreberiana
Russia: Murmansk Prov-
ince
M-M-1569 (MW)
MW881239
-
MW881242
-
OQ199085
RF41
Dicranella cf. schreberiana
Russia: Altai Mts.
MW9031017
MN178090
-
MN187509
-
MN178016
FDt150
Dicranella schreberiana
Russia: Moscow
MW9111436
OQ199126
-
OQ199126
-
OQ199087
TJH17
Dicranella schreberiana
Netherlands
Aptroot 69819 (L)
MN178089
-
MN187508
-
MN178015
TJH16
Dicranella schreberiana var.
robusta
Netherlands
Nieuwkoop 2012060 (L)
MN178091
-
MN187510
-
MN178017
TJH23
Dicranella schreberiana var.
robusta
Norway
Siebel 2014.732 (L)
MN178092
-
MN187511
-
MN178018
TJH35
Dicranella schreberiana var.
robusta
Netherlands
Siebel 2015.561 (L)
MN178094
-
MN187513
-
MN178019
FDt151
Dicranella schreberiana var.
robusta
Russia: Moscow Province
MW9031001
OQ199125
-
OQ199125
-
OQ199086
MDP300
Hygrodicranum bolivianum
Buck, 39497 (DUKE)
-
-
AY908115
-
AY908904
IPG20
Hygrodicranum herrerae
Chile
Stech 15-028 (L)
MN178113
-
MN187531
-
MN178037
MDP488
Polymerodon andinus
M. Lewis 87608 (DUKE)
-
-
AY908166
-
AY908903
RF66
Dicranella hookeri
Greene 2988 (LE)
MW881240
-
MW881243
-
-
RF65
Dicranella hookeri
Chile
Greene 1265 (LE)
-
-
OQ199127
-
-

Plants 2023, 12, 1360 37 of 40
FDt198
Dicranella hookeri
Chile
Larraín 38481 (MW)
OQ199128
-
OQ199128
-
OQ199088
TJH13
Dicranella campylophylla
Chile
Stech 15-006 (L)
MN178062
-
MN187482
-
MN177993
TJH04
Dicranella campylophylla
Chile
Stech 15-007 (L)
MN178061
-
MN187481
-
MN177992
RF67
Dicranella campylophylla
R. Smith 2763 (LE)
MW881241
-
MW881244
-
-
FDt196
Dicranella campylophylla
Chile
Larraín 45977 (MW)
OQ199129
-
OQ199129
-
OQ199089

Plants 2023, 12, 1360 38 of 40
References
1. Ignatov, M.S.; Huttunen, S. Brachytheciaceae (Bryophyta)—A family of sibling genera. Arctoa 2002, 11, 245–296.
https://doi.org/10.15298/arctoa.11.20.
2. Vanderpoorten, A.; Hedenäs, L.; Cox, C.J.; Shaw, A.J. Phylogeny and Morphological Evolution of the Amblystegiaceae
(Bryopsida). Mol. Phylogenetics Evol. 2002, 23, 1–21. https://doi.org/10.1006/mpev.2001.1067.
3. Vanderpoorten, A.; Hedenäs, L.; Cox, C.J.; Shaw, A.J. Circumscription, classification, and (Bryopsida) inferred from nuclear and
chloroplast DNA sequence data and morphology taxonomy of Amblystegiaceae. Taxon 2002, 51, 115–122.
https://doi.org/10.2307/1554968.
4. Hedenäs, L. Additional insights into the phylogeny of Calliergon, Loeskypnum, Straminergon, and Warnstorfia (Bryophyta:
Calliergonaceae). J. Hattori Bot. Lab. 2006, 100, 125–134.
5. Olsson, S.; Buchbender, V.; Enroth, J.; Hedenäs, L.; Huttunen, S.; Quandt, D. Phylogenetic analyses reveal high levels of
polyphyly among pleurocarpous lineages as well as novel clades. Bryologist 2009, 112, 447–466. https://doi.org/10.1639/0007-
2745-112.3.447.
6. Plášek, V.; Sawicki, J.; Ochyra, R.; Szczecińska, M.; Kulik, T. New taxonomical arrangement of the traditionally conceived genera
Orthotrichum and Ulota (Orthotrichaceae, Bryophyta). Acta Musei Sil. Sci. Nat. 2015, 64, 169–174.
7. Kučera, J.; Kuznetsova, O.I.; Manukjanová, A.; Ignatov, M.S. A phylogenetic revision of the genus Hypnum: Towards
completion. Taxon 2019, 68, 628–660. https://doi.org/10.1002/tax.12095.
8. Fedosov, V.E.; Fedorova, A.V.; Fedosov, A.E.; Ignatov, M.S. Phylogenetic inference and peristome evolution in haplolepideous
mosses, focusing on Pseudoditrichaceae and Ditrichaceae s.l. Bot. J. Linn. Soc. 2016, 181, 139–155.
https://doi.org/10.1111/boj.12408.
9. Fedosov, V.E.; Fedorova, A.V.; Larraín, J.; Bonfim-Santos, M.; Stech, M.; Kučera, J.; Brinda, J.C.; Tubanova, D.Y.; von Konrat,
M.; Ignatova, E.A.; et al. Unity in diversity: Phylogeny and taxonomy of Rhabdoweisiaceae (Dicranales, Bryophyta). Bot. J. Linn.
Soc. 2021, 195, 545–567.
10. Santos, M.B.; Fedosov, V.; Hartman, T.; Fedorova, A.; Siebel, H.; Stech, M. Phylogenetic inferences reveal deep polyphyly of
Aongstroemiaceae and Dicranellaceae within the haplolepideous mosses (Dicranidae, Bryophyta). Taxon 2021, 70, 246–262.
https://doi.org/10.1002/tax.12439.
11. Medina, R.; Johnson, M.; Liu, Y.; Wilding, N.; Hedderson, T.A.; Wickett, N.; Goffinet, B. Evolutionary dynamism in bryophytes:
Phylogenomic inferences confirm rapid radiation in the moss family Funariaceae. Mol. Phylogenetics Evol. 2018, 120, 240–247.
https://doi.org/10.1016/j.ympev.2017.12.002.
12. Medina, R.; Johnson, M.G.; Liu, Y.; Wickett, N.; Shaw, A.J.; Goffinet, B. Phylogenomic delineation of Physcomitrium (Bryophyta:
Funariaceae) based on targeted sequencing of nuclear exons and their flanking regions rejects the retention of Physcomitrella,
Physcomitridium and Aphanorrhegma. J. Syst. Evol. 2019, 57, 404–417.
13. Ignatov, M.S.; Ignatova, E.A. Mosses of Middle Part of European Russia; KMK Press: Moscow, Russia, 2004; Volume 2, pp. 609–
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