Journal of Bryology
ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/yjbr20
An annotated checklist of bryophytes of Europe,
Macaronesia and Cyprus
N. G. Hodgetts , L. Söderström , T. L. Blockeel , S. Caspari , M. S. Ignatov ,
N. A. Konstantinova , N. Lockhart , B. Papp , C. Schröck , M. Sim-Sim , D.
Bell , N. E. Bell , H. H. Blom , M. A. Bruggeman-Nannenga , M. Brugués , J.
Enroth , K. I. Flatberg , R. Garilleti , L. Hedenäs , D. T. Holyoak , V. Hugonnot ,
I. Kariyawasam , H. Köckinger , J. Kučera , F. Lara & R. D. Porley
To cite this article: N. G. Hodgetts , L. Söderström , T. L. Blockeel , S. Caspari , M. S. Ignatov ,
N. A. Konstantinova , N. Lockhart , B. Papp , C. Schröck , M. Sim-Sim , D. Bell , N. E. Bell , H.
H. Blom , M. A. Bruggeman-Nannenga , M. Brugués , J. Enroth , K. I. Flatberg , R. Garilleti , L.
Hedenäs , D. T. Holyoak , V. Hugonnot , I. Kariyawasam , H. Köckinger , J. Kučera , F. Lara & R. D.
Porley (2020) An annotated checklist of bryophytes of Europe, Macaronesia and Cyprus, Journal of
Bryology, 42:1, 1-116, DOI: 10.1080/03736687.2019.1694329
To link to this article: https://doi.org/10.1080/03736687.2019.1694329
© 2020 The Author(s). Published by Informa
UK Limited, trading as Taylor & Francis
Group
Published online: 28 May 2020.
Submit your article to this journal
Article views: 921
View related articles
View Crossmark data
Citing articles: 1 View citing articles
Full Terms & Conditions of access and use can be found at
https://www.tandfonline.com/action/journalInformation?journalCode=yjbr20
JOURNAL OF BRYOLOGY
2020, VOL. 42, NO. 1, 1–116
https://doi.org/10.1080/03736687.2019.1694329
BRYOLOGICAL MONOGRAPH
An annotated checklist of bryophytes of Europe, Macaronesia and Cyprus
N. G. Hodgettsa, L. Söderström b, T. L. Blockeelc, S. Casparid, M. S. Ignatove, N. A. Konstantinovaf,
N. Lockhartg, B. Papph, C. Schröcki, M. Sim-Simj, D. Bellk, N. E. Bell k, H. H. Bloml, M. A. BruggemanNannengam, M. Bruguésn, J. Enrotho, K. I. Flatbergp, R. Garilleti q, L. Hedenäs r, D. T. Holyoaks,
V. Hugonnott, I. Kariyawasamk, H. Köckingeru, J. Kučera v, F. Lara w and R. D. Porleyx
a
Cuillin Views, Isle of Skye, UK; bNorwegian University of Science and Technology, Trondheim, Norway; cDore, Sheffield, UK; dZentrum für
Biodokumentation, Schiffweiler, Saarland, Germany; eMoscow University and Tsitsin Main Botanica Garden RAS, Moscow, Russia; fPolarAlpine Botanical Garden-Institute RAS, Kirovsk, Russia; gNational Parks and Wildlife Service, Dublin, Ireland; hHungarian Natural History
Museum, Budapest, Hungary; iBiology Centre of Upper Austrian National Museums, Linz, Austria; jUniversidade de Lisboa, Faculdade de
Ciências, Departamento de Biologia Vegetal, cE3c - Centre for Ecology, Evolution and Environmental Changes, Lisboa, Portugal; kRoyal
Botanic Garden, Edinburgh, UK; lNorwegian Institute of Bioeconomy Research, Bergen, Norway; mGriffensteijnseplein 23, Zeist, The
Netherlands; nFacultat de Biosciènces, Universitat Autònoma de Barcelona, Spain; oOrganismal and Evolutionary Research Programme,
University of Helsinki, Finland; pDepartment of Natural History, University Museum, Trondheim, Norway; qDepartamento de Botánica y
Geología, Facultad de Farmacia, Universidad de Valencia, Spain; rSwedish Museum of Natural History, Stockholm, Sweden; sRua da Barcoila
274, Cabeçudo, Portugal; tConservatoire Botanique National du Massif Central, Chavaniac Lafayette, France; uRoseggergasse 12,
Weisskirchen, Austria; vFaculty of Science, University of South Bohemia, České Budějovice, Czech Republic; wCentro de Investigación en
Biodiversidad y Cambio Global (CIBC-UAM), Madrid, Spain; xCerca dos Pomares, Aljezur, Portugal
ABSTRACT
ARTICLE HISTORY
Introduction. Following on from work on the European bryophyte Red List, the
taxonomically and nomenclaturally updated spreadsheets used for that project have been
expanded into a new checklist for the bryophytes of Europe.
Methods. A steering group of ten European bryologists was convened, and over the course
of a year, the spreadsheets were compared with previous European checklists, and all
changes noted. Recent literature was searched extensively. A taxonomic system was
agreed, and the advice and expertise of many European bryologists sought.
Key results. A new European checklist of bryophytes, comprising hornworts, liverworts and
mosses, is presented. Fifteen new combinations are proposed.
Conclusions. This checklist provides a snapshot of the current European bryophyte flora
in 2019. It will already be out-of-date on publication, and further research, particularly
molecular work, can be expected to result in many more changes over the next few years.
First Published Online
12 May 2020
KEYWORDS
Bryophytes; mosses;
liverworts; hornworts;
Europe; checklist
Introduction
The last published annotated checklist for mosses in
Europe was Hill et al. (2006), while for liverworts and
hornworts it was Grolle and Long (2000), although a
checklist with distributional records was produced by
Söderström, Urmi, et al. (2002) and updated by
Söderström, Urmi, et al. (2007). In addition, Séneca
and Söderström (2009) published a checklist of the
Sphagnophyta of Europe and Macaronesia with distribution data. During work for the new European
bryophyte Red List, it became apparent that not
only was it necessary to publish a new European
checklist covering all the bryophytes, but also that
this would be relatively little extra work, because
spreadsheets of species indicating their distribution
by country had been prepared for the Red List, and
every effort made to update the nomenclature and
taxonomy. A new checklist is also needed because
of the great amount of new molecular work that
CONTACT N. G. Hodgetts
has been done (and continues to be done) on bryophytes, revealing many new and often unexpected
relationships. The nomenclature of liverworts and
hornworts in particular has undergone radical
changes, as recorded in the recent World Checklist
(Söderström, Hagborg, et al. 2016), which, however,
did not contain synonyms.
Ironically, the checklist by Hill et al. (2006) was prepared in response to the intention of the European
Committee for the Conservation of Bryophytes
(ECCB) to produce a new Red List, but the latter
had to wait another ten years before being
addressed properly, when the ECCB entered into collaboration with the International Union for the Conservation of Nature (IUCN), at which point LIFE
funding became available for the Red List project.
The Red List has now been published (Hodgetts
et al. 2019), and the names are substantially the
nick1901@hotmail.co.uk
© 2020 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (http://creativecommons.org/licenses/bync-nd/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited, and is not altered, transformed, or
built upon in any way.
Published online 28 May 2020
2
N. G. HODGETTS ET AL.
same as in this checklist, although there have been
some updates and additions to the checklist since
the Red List cut-off date of 31 December 2018, and
the geographical areas covered by the Red List and
the checklist differ slightly (see below).
the World Checklist (Söderström, Hagborg, et al.
2016). The provisional spreadsheets were also sent
to other bryologists for further expert advice
(authors DB, VH and HK and those listed in acknowledgements below).
Methods
Taxonomic hierarchy
Following work on the Red List, funding was secured
from the Irish government’s National Parks and Wildlife Service to enable author NGH to co-ordinate a
new checklist project. A Steering Group was set
up consisting of Tom Blockeel, Steffen Caspari, Nick
Hodgetts, Misha Ignatov, Nadya Konstantinova, Neil
Lockhart, Beata Papp, Christian Schröck, Manuela
Sim-Sim and Lars Söderström. Communication has
largely been through e-mail, but the Steering
Group met once in Budapest, in November 2018,
to coincide with the final Red List meeting. After
this meeting, NGH led on the moss part of the
checklist and author LS led on the liverworts and
hornworts.
The existing network of ECCB contacts was crucial to
the project, and a large number of bryologists in
Europe have contributed, and those making major contributions are included in the authorship.
The first step was to compare the data in the Red List
spreadsheets, which had been kept up-to-date taxonomically and nomenclaturally, with that in Hill et al.
(2006) and Grolle and Long (2000). Differences
between these checklists and current concepts were
highlighted and marked for comment. These spreadsheets were then examined closely, species by
species, at the Steering Group meeting, with two subgroups convened, one for mosses, the other for liverworts and hornworts. A team of experts on particular
taxonomic groups of mosses was recruited to advise
on their areas of expertise. These are listed below
and are included in the authorship of this paper:
For the liverworts and hornworts, the taxonomic hierarchy follows the Catalogue of Life (Roskov et al.
2019), the online database of the world’s known
species of animals, plants, fungi and micro-organisms, in which the hierarchy for the hornworts
(Anthocerotophyta) and the liverworts (Marchantiophyta) is contributed by the Early Land Plants
Today project (Söderström, Hagborg, et al. 2019).
The higher classification is substantially the same as
the one presented in the bryophyte phylogeny
poster by Cole et al. (2019).
It was more complicated to agree on a higher
classification of mosses, as current concepts are in
a considerable state of flux as more molecular
work is done. Pleurocarpous mosses in particular
have undergone many changes in recent years,
and it is expected that this will continue before relative stability is achieved. Our starting point was Frey
and Stech (2009), but it has been necessary to
update this treatment with the results of new
research. The resultant higher classification is substantially in agreement with the poster produced
recently by Cole et al. (2019), which is probably the
most accurate representation of the higher classification of mosses as it is currently understood,
although there are some differences. For example,
Cole et al. (2019) place Hypopterygium in the order
Hypopterygiales, while we have it in the Hypopterygiaceae within the order Hookeriales.
The final higher classification presented in this
checklist is not offered as a definitive view, and it can
be expected that there will be further changes with
more research.
Neil Bell, Isuru Kariyawasam – Polytrichaceae
Hans Blom – Schistidium
Ida Bruggeman-Nannenga – Fissidentaceae
Monserrat Brugués – Funariaceae
Johannes Enroth – Neckeraceae
Kjell Ivar Flatberg – Sphagnaceae
Lars Hedenäs – Amblystegiaceae, Calliergonaceae,
pleurocarps
David Holyoak – Bryaceae
Jan Kučera – Pottiaceae
Paco Lara, Ricardo Garilleti – Orthotrichaceae
Ron Porley – Grimmia and Coscinodon
Specialist advice was either not needed or not available for other groups. It was less crucial to have
specialist advice on taxonomic groups of liverworts,
as much of the work had been done previously for
List of taxa
As Hill et al. (2006) and Grolle and Long (2000) were
the starting points for this checklist, taxa that have
remained the same since these publications are
mostly listed without further comment. All synonyms
that have been used since these publications are
included. In addition, some synonyms whose synonymy predates these publications, but have continued to be in common usage, have been included.
The liverworts in particular have undergone some
radical taxonomic and nomenclatural changes since
Grolle and Long (2000). Most of these are recorded,
but without synonyms, in the World Checklist (Söderström, Hagborg, et al. 2016). The synonymy for the
JOURNAL OF BRYOLOGY
liverworts is therefore considerably more extensive
than that for the mosses.
Hybrid taxa are mentioned only if they have been
treated in recent literature.
Author citations
We have tried to follow the International Plant
Names Index (www.ipni.org) for author citations,
although no doubt we will not always have succeeded. In practice, most author citations, except
for very recently treated taxa, are the same as
in Hill et al. (2006) for the mosses and Söderström,
Hagborg, et al. (2016) for the liverworts and
hornworts, but a small number of citations have
been corrected.
Bryum s.lat.
It is necessary to make some remarks about Bryum
s.lat. as the current position is confusing and unsatisfactory. The reviews of world-wide scope by Ochi
(1959, 1970, 1972, 1980, 1981, etc.) placed all European Bryaceae in genus Bryum with several subgenera, on the basis of extensive morphological
studies. Other workers treated his subgenera as
genera, Anomobryum, Brachymenium, Bryum, Plagiobryum, Rhodobryum (e.g. Holyoak in Hill et al. 2006,
Guerra, Gallego, et al. 2010). Molecular data (Pedersen and Hedenäs 2002, 2003; Pedersen, Cox and
Hedenäs 2003; Pedersen, Holyoak and Newton
2007) showed this generic treatment to be illfounded, since several of the widely recognised subgenera/genera were shown to be polyphyletic. It
also became apparent that morphology in Bryaceae
often provides entirely misleading evidence of phylogeny: Examples of this include the morphologically
striking ‘Plagiobryum’ zieri nested within a clade containing e.g. Ptychostomum pseudotriquetrum, the
superficially similar small tuber-bearing ‘Erythrocarpa’ belonging in at least two distinct genera,
and some Bryum dichotomum being difficult to separate morphologically from Ptychostomum imbricatulum (syn. Bryum caespiticium). Thus, morphological
evolution in Bryaceae appears to be ‘decoupled’
from phylogeny (Holyoak and Pedersen 2007).
Generic allocations based on morphology alone are
therefore prone to error, some through convergent
evolution of distant relatives, others by rapid
change among close relatives. These findings have
not been refuted by subsequent studies and parts
of the sequence data used have been confirmed
and supplemented by other workers (Guerra,
Jiménez-Martínez, Cano, Brugués 2008; Guerra,
Jiménez-Martínez, Cano, Jiménez-Fernández 2011;
Wang and Zhao 2009; Bell, Long and Hollingsworth
3
2013). Thus the generic taxonomy adopted here is
based mainly on molecular data.
Nevertheless, Spence (2014) and e.g. Ochyra, Plášek,
et al. (2018) have largely ignored the implications of
the molecular results and used morphology almost
exclusively for their generic classifications. Parts of the
classifications by Spence are directly contradicted by
molecular data (cf. Shaw 2014: p.10–11); much of the
remainder is at risk of being erroneous due to convergent evolution.
It is frustrating that a lack of further molecular
data still prevents the allocation of some species to
segregate genera, but this problem is declining as
more species are sequenced. Indeed, some unpublished DNA barcoding and nuclear genomic sequencing data from one of the authors (DB), along with
some last-minute sequencing by another (MSI), and
interpretation by a third (DTH), have resulted in a
clarification of generic affinity for several species.
Further sequencing work should see significant
increases in the data available within the next few
years. The alternative of reverting to an enormous
and still partly polyphyletic genus Bryum thus
seems to be a poorer option. It would also be out
of step with the rapidly improving classifications
of some other moss families. Currently recognised
European genera of Bryaceae in this checklist are
Anomobryum, Brachymenium, Bryum, Imbribryum,
Ptychostomum and Rhodobryum. We do not recognise Gemmabryum, Osculatia, Plagiobryoides, Plagiobryum or Rosulabryum.
Area covered
For the purposes of this checklist, Europe is regarded in
the same way as it was in Hill et al. (2006), but with the
addition of Cyprus. Thus, the following territories at the
margins of Europe are included:
All of Russia west of the Ural watershed and to the
north of the Caucasus watershed;
Western Kazakhstan;
Iceland;
Svalbard;
Jan Mayen;
Novaya Zemlya;
Franz Josef Land;
Malta;
all the Greek Islands;
all other Mediterranean islands belonging to European
countries (e.g. Pantelleria);
Turkey west of the Bosporus (i.e. Turkey-in-Europe);
Azores;
Canary Islands;
Madeira.
4
N. G. HODGETTS ET AL.
Table 1. Bryophytes introduced to Europe
Species
Location
Native range
Liverworts
Heteroscyphus fissistipus
Lophocolea bispinosa
Lophocolea brookwoodiana
Lophocolea semiteres
Sphaerocarpos stipitatus
Tricholepidozia lindenbergii var. lindenbergii
Tricholepidozia tetradactyla
garden in Ireland
gardens and naturalised in Britain, Ireland & Spain
cemetery in southern England
naturalised in western Europe
Portugal
gardens in Wales
gardens in Britain
Australia, New Zealand
Australia
Unknown
Southern Hemisphere
Southern Hemisphere
New Zealand, Philippines and Fiji
New Zealand
Mosses
Achrophyllum dentatum
Atrichum crispum
Calomnion complanatum
Calyptrochaeta apiculata
Campylopus introflexus
Dicranoloma menziesii
Hennediella macrophylla
Hennediella stanfordensis
Hypopterygium tamarisci
Leptodontium proliferum
Leptotheca gaudichaudii
Myuroclada maximowiczii
Orthodontium lineare
Sematophyllum adnatum
Thuidiopsis sparsa
Tortula amplexa
garden in England
naturalised in Britain & Ireland
gardens in Ireland
naturalised in southern England & Ireland
naturalised & widespread in much of Europe
on tree ferns in garden in Ireland
naturalised in Britain
naturalised in Britain, Ireland, France & Greece
gardens in Portugal & Italy
bowling green in northern England
gardens in southern England & Ireland
garden in southern Russia
naturalised & widespread in western and central Europe
naturalised in northern Italy, Switzerland & Hungary
former park in Madeira
naturalised in England
Southern Hemisphere
Eastern North America
Australia, New Zealand
Southern Hemisphere
Southern Hemisphere
Southern Hemisphere
New Zealand
California, Mexico & Australia
Tropical Africa & America
Southern Hemisphere
Southern Hemisphere
Eastern Asia, Alaska
Southern Hemisphere
America
Southern Hemisphere
Western North America
The decision to include Cyprus was made because it is
politically part of Europe, although geographically
closer to Asia. In practice, its inclusion or exclusion
is fairly academic, as there are no bryophyte taxa
known that have their only ‘European’ occurrence
in Cyprus, except for the dubious species Funaria
anomala, and possibly Grimmia ungeri, which has
been considered to be endemic to Cyprus by
some authors.
The area covered differs slightly from that covered
by the Red List (IUCN 2019), which excludes the Caucasus. Both the checklist and the Red List exclude Cape
Verde (Cabo Verde).
Composition of the bryophyte flora
The moss flora of Europe, as described in this checklist, comprises 1390 species, plus 45 subspecies and
104 varieties. There are 494 species of liverworts,
with 35 subspecies and 56 varieties, and 8 species
of hornworts, with no subspecies or varieties,
giving a total of 1892 species of bryophytes in
Europe, plus 80 subspecies and 160 varieties. These
figures only include subspecies and varieties where
species are represented by more than one subspecies or variety in Europe. There are 187 endemic
taxa, with 56 of these being endemic to the Macaronesian islands. The level at which a taxon is recognised is always to some extent arbitrary, even with
the benefit of molecular data. In general, we have
included subspecies and varieties if they have been
validly published and there is no good reason to
reject them. However, this self-imposed guidance
has been considerably modified according to
expert opinion from many of the authors and those
listed in the acknowledgements. It is indicated in
the footnotes where disagreement exists.
There are 23 obvious introductions that are now
more or less naturalised in at least some part of
Europe. These are listed in Table 1. Taxa are considered
introductions only if they have clearly been introduced
by human agency. Other species that might be
regarded as introductions, that may for example be
chance ephemeral colonists from wind-blown spores,
are regarded as native, as they apparently arrived
through natural agency. Whether any particular taxon
is native or not to Europe is of course arguable. To
some extent the native status of taxa matters less for
bryophytes than it does for vascular plants, since
their often very efficient dispersal mechanisms mean
that many species can potentially occur in suitable
habitats worldwide. Several further species were
listed by Hill et al. (2006) as non-native (Anomobryum
apiculatum, Bryum valparaisense, Chenia leptophylla,
Splachnobryum obtusum, Syntrichia bogotensis, Tortula
bogosica, Tortula bolanderi), but there is significant
doubt about their non-native status, and they are not
listed in Table 1.
New validations
Three liverwort taxa were published invalidly and
therefore require validation.
Riccia sorocarpa subsp. arctica R.M.Schust. ex Köckinger & L.Söderstr., subsp. nov. [Riccia sorocarpa var.
arctica R.M.Schust., J. Hattori Bot. Lab. 71: 274, 1992,
nom. inval. ICN Art. 37.7]. Holotype: Greenland, Kangerdlugssuak, Inglefield Bay, NW. Greenland, R.M.Schuster
JOURNAL OF BRYOLOGY
45831 (F-C0000489F; http://emuweb.fieldmuseum.org/
botany/Display.php?irn=9894&QueryPage=%2Fbotany
%2Fdetailed.php). The description in Schuster (1992a)
together with the current data on type specimen
validates the name. This subspecies was described
from Greenland but also occurs in the Alps
(Köckinger 2017).
Riccia sorocarpa subsp. erythrophora R.M.Schust. ex
Konstant. & L.Söderstr., subsp. nov. [Riccia sorocarpa
subsp. erythrophora R.M.Schust., J. Hattori Bot. Lab.
71: 274, 1992, nom. inval. ICBN Art. 37.7]. Holotype: California: 1.5 km W of Potrero, San Diego Co., 1982,
R.M.Schuster 82-146 (F-C1049209F; http://emuweb.
fieldmuseum.org/botany/Display.php?irn=2346544&
QueryPage=%2Fbotany%2Fdetailed.php; type originally filed under Riccia nigrella). The description
in Schuster (1992a) together with the current
data on the type specimen validate the name. It was
described from California but is also known from
southern Russia (Konstantinova and Doroshina 2011).
Scapania paludicola var. rotundiloba R.M.Schust. ex
Konstant. & L.Söderstr., var. nov. Holotype: “West
Greenland: east end of Nûk Peninsula, Alángup nunâ,
68°42’N., 52°10’W. Schuster & Damsholt RMS 70-2878”
(F-C0171445F;
http://emuweb.fieldmuseum.org/
botany/Display.php?irn=2145467&QueryPage=%2F
botany%2Fdetailed.php) [Scapania paludosa var.
rotundiloba R.M.Schust., Hepat. Anthocerotae N. Amer.
3: 519, 1974, nom. inval. ICBN Art. 37.2; based on more
than one gathering]. The description in Schuster (1974:
519), together with the type selected here, validates
the name.
New combinations
New combinations are made by individual authors.
Mesoptychia gillmanii var. laxa (Schiffn. ex Burrell)
L.Söderstr. comb. nov. Basionym: Lophozia schultzii
var. laxa Schiffn. ex Burrell, J. Bot. 49: 217, 1911 (see
Burrell 1911).
Anomobryum apiculatum (Schwägr.) D.Bell & Holyoak
comb. nov. Basionym: Bryum apiculatum Schwägr., Sp.
Musc. Frond., Suppl. 1, sect. 2, p. 102, pl. 72 [top], 1816
(see Schwägrichen 1816).
Anomobryum notarisii (Mitt.) D.Bell & Holyoak comb.
nov. Basionym: Bryum notarisii Mitt., J. Linn. Soc., Bot.,
7, p. 3, 1864 (see Mitten 1864).
Didymodon glaucus subsp. verbanus (W.E.Nicholson &
Dixon) Jan Kučera comb. nov. Basionym: Eucladium
5
verbanum W.E.Nicholson & Dixon, Revue Bryologique
39: 89. 1912 (see Dixon 1912).
Fissidens
subgenus
Pachyfissidens
(Müll.Hal.)
L.Söderstr. & A.Hagborg, comb. nov. Basionym: Fissidens sect. Pachyfissidens Müll.Hal., Syn. Musc. Frond.
1: 45, 1849 [1848] (see Müller 1849). Note: The subgenus has frequently been cited from Kindberg (1897),
but in that paper Pachyfissidens is an unranked
taxon. We have not been able to localise any valid
combination at subgeneric rank and thus propose
it here.
Imbribryum subapiculatum (Hampe) D.Bell &
Holyoak comb. nov. Basionym: Bryum subapiculatum Hampe, Vidensk. Meddel. Dansk Naturhist.
Foren. Kjøbenhavn, ser. 3, 4, p. 51, 1872 (see Hampe
1872).
Imbribryum tenuisetum (Limpr.) D.Bell & Holyoak
comb. nov. Basionym: Bryum tenuisetum Limpr., Jahresber. Schles. Ges. Vaterl. Cult., 74(2b), p. 4, 1897 (see Limpricht 1897).
Mnioideae L.Söderstr., N.G.Hodgetts & Ignatov, comb.
et stat. nov.
Basionym: Trib. Mnieae Müll., Syn. Musc. Frond.: 152,
1858 (Müller 1858) ‘Mnioideae’.
≡ Mnioideae A.J.E.Sm., Moss Flora Brit. Ireland, 2nd ed.:
615, 2004 (Smith 2004), nom. inval. (ICN Art. 39.1; no
Latin description).
≡ Mnioideae Stech & W.Frey, Syll. Pl. Fam. 3: 196 (Frey
& Stech 2009), nom. inval. (ICN Art. 39.1; no Latin
description).
≡ Mnioideae T.J.Kop., Acta Bryolich. Asiatica 5: 39
(Koponen 2014), nom. inval. (ICN Art. 38.1(a); no
description).
Type genus: Mnium Hedw.
Note: The subfamily Mnioideae has been recognized
for a long time but as far as we know never been validated. Most authors seem to assume it is an autonym
under Mniaceae (e.g. Smith 2004; Frye and Stech
2009). However, autonyms only exist under genera
and species (ICN Art. 22.1 and 26.1; Turland et al.
2018) so a formal description of the taxon is
needed. No description before 2011 is in Latin, as
required then, or refers to Müller (1858) to make it
a valid combination. Latin descriptions have been
unnecessary since 2011, but we have not found any
recent description of any sort that fulfils all the criteria. Frey and Stech (2009) give a good description,
but they seem to treat all subfamilies based on the
family name as autonyms, contrary to the code
(Turland et al. 2018). A reference to Müller (1858),
who treats the taxon as a tribus (although with an
6
N. G. HODGETTS ET AL.
incorrect ending according to the current code), validates the name.
Platyhypnum tatrense (Váňa) Hedenäs & Ignatov
comb. nov. Basionym: Ochyraea tatrensis Váňa,
J. Bryol. 14: 261. f. 1-3. 1986 (see Váňa 1986).
Ptychostomum austriacum (Köckinger, Holyoak &
Suanjak) D.Bell & Holyoak comb. nov. Basionym:
Bryum austriacum Köckinger, Holyoak & Suanjak,
J. Bryol. 35(1), p. 57, 2013 (see Köckinger, Holyoak,
et al. 2013).
Ptychostomum cellulare (Hook.) D.Bell & Holyoak comb.
nov. Basionym: Bryum cellulare Hook. in Schwägr., Sp.
Musc. Frond., Suppl. 3, vol. 1, sect. 1, p. 9, pl. 214a, fig.
a, 1827 (see Schwägrichen 1827).
Ptychostomum elegans (Nees.) D.Bell & Holyoak comb.
nov. Basionym: Bryum elegans Nees, in Bridel, Bryol.
Univ. 1 (2): p. 849. 1827 (see Bridel 1827).
Ptychostomum minii (Podp. ex Guim.) D.Bell & Holyoak
comb. nov. Basionym: Bryum minii Podp. ex Guim., Rev.
Bryol. Lichénol., 8, p. 112–114, 1935.
Ptychostomum schleicheri (DC.) J.R.Spence ex D.Bell
& Holyoak comb. nov. Basionym: Bryum schleicheri
DC. in Lam., Fl. Franç., 6, p. 226, 1815 (see Lamarck
1815).
Ptychostomum schleicheri var. latifolium (Schwägr.)
D.Bell & Holyoak comb. nov. Basionym: Mnium
latifolium Schwägr., Spec. Musc. Suppl. 1(2), p. 138,
1816 ≡ Bryum schleicheri var. latifolium (Schwägr.)
Schimp.
Taxonomic synopsis
This synopsis lists the higher taxa relevant to Europe
down to family level.
Anthocerotophyta Stotler & Crand.-Stotl.
Anthocerotopsida de Bary ex Jancz.
Anthocerotidae Rosenv.
Anthocerotales Limpr.
Anthocerotaceae Dumort.
Phymatocerotales R.J.Duff
Phymatocerotaceae R.J.Duff
Notothylatidae R.J.Duff
Notothyladales Hyvönen & Piippo
Notothyladaceae Müll.Frib. ex Prosk.
Phaeocerotideae Hässel
Marchantiophyta Stotler & Crand.-Stotl.
Haplomitriopsida Stotler & Crand.-Stotl.
Haplomitriidae Stotler & Crand.-Stotl.
Calobryales Hamlin
Haplomitriaceae Dědeček
Jungermanniopsida Stotler & Crand.-Stotl.
Jungermanniidae Engl.
Jungermanniales H.Klinggr.
Cephaloziineae Schljakov.
Adelanthaceae Grolle
Anastrophyllaceae L.Söderstr., De Roo &
Hedd.
Cephaloziaceae Mig.
Cephaloziellaceae Douin
Lophoziaceae Cavers
Scapaniaceae Mig.
Jungermanniineae R.M.Schust. ex Stotler &
Crand.-Stotl.
Acrobolbaceae E.A.Hodgs.
Antheliaceae R.M.Schust.
Arnelliaceae Nakai
Calypogeiaceae Arnell
Endogemmataceae Konstant.
Geocalycaceae H.Klinggr.
Gymnomitriaceae H.Klinggr.
Harpanthaceae Arnell
Hygrobiellaceae Konstant. & Vilnet
Jungermanniaceae Rchb.
Saccogynaceae Heeg
Solenostomataceae Stotler & Crand.-Stotl.
Southbyaceae Váňa, Crand.-Stotl., Stotler &
D.G.Long
Lophocoleineae Schljakov
Blepharostomataceae W.Frey & M.Stech
Herbertaceae Müll.Frib. ex Fulford & Hatcher
Lepidoziaceae Limpr.
Lophocoleaceae Vanden Berghen
Mastigophoraceae R.M.Schust.
Plagiochilaceae Müll.Frib.
Trichocoleaceae Nakai
Myliineae J.J.Engel & Braggins ex Crand.-Stotl.
Myliaceae Schljakov
Porellales Schljakov
Jubulineae Müll.Frib.
Frullaniaceae Lorch
Jubulaceae H.Klinggr.
Lejeuneaceae Cavers
Porellineae R.M.Schust.
Porellaceae Cavers
Radulineae R.M.Schust.
Radulaceae Müll.Frib.
Ptilidiales Schljakov
Ptilidiaceae H.Klinggr.
Metzgeriidae Barthol.-Began
Metzgeriales Chalaud
Aneuraceae H.Klinggr.
Metzgeriaceae H.Klinggr.
Pleuroziales Schljakov
Pleuroziaceae Müll.Frib.
Pelliidae He-Nygrén
Fossombroniales Schljakov
JOURNAL OF BRYOLOGY
Calyculariineae He-Nygrén, Juslén, Ahonen,
Glenny & Piippo
Calyculariaceae
He-Nygrén,
Juslén,
Ahonen, Glenny & Piippo
Fossombroniineae R.M.Schust. ex Stotler &
Crand.-Stotl.
Fossombroniaceae Hazsl.
Petalophyllaceae Stotler & Crand.-Stotl.
Pallaviciniales W.Frey & M.Stech
Pallaviciniineae R.M.Schust.
Moerckiaceae K.I.Goebel ex Stotler &
Crand.-Stotl.
Pallaviciniaceae Mig.
Pelliales He-Nygrén
Pelliaceae H.Klinggr.
Marchantiopsida Cronquist, Takht. & W.Zimm.
Blasiidae He-Nygrén
Blasiales Stotler & Crand.-Stotl.
Blasiaceae H.Klinggr.
Marchantiidae Engl.
Lunulariales H.Klinggr.
Lunulariaceae H.Klinggr.
Marchantiales Limpr.
Aytoniaceae Cavers
Cleveaceae Cavers
Conocephalaceae Müll.Frib. ex Grolle
Corsiniaceae Engl.
Cyathodiaceae Stotler & Crand.-Stotl.
Dumortieraceae D.G.Long
Exormothecaceae Müll.Frib. ex Grolle
Marchantiaceae Lindl.
Oxymitraceae Müll.Frib. ex Grolle
Ricciaceae Rchb.
Targioniaceae Dumort.
Sphaerocarpales Cavers
Riellaceae Engl.
Sphaerocarpaceae Heeg
Bryophyta Schimp.
Sphagnophytina Doweld
Sphagnopsida Schimp.
Sphagnales Limpr.
Sphagnaceae Dumort.
Bryophytina Engler
Andreaeopsida J.H.Schaffn.
Andreaeidae Engl.
Andreaeales Limpr.
Andreaeaceae Dumort.
Oedipodiopsida Goffinet & W.R.Buck
Oedipodiales Goffinet & W.R.Buck
Oedipodiaceae Schimp.
Tetraphidopsida Goffinet & W.R.Buck
Tetraphidales M.Fleisch.
Tetraphidaceae Schimp.
Polytrichopsida Doweld
Polytrichales M.Fleisch.
Polytrichaceae Schwägr.
Bryopsida Pax
7
Buxbaumiidae Doweld
Buxbaumiales M.Fleisch.
Buxbaumiaceae Schimp.
Diphysciidae Ochyra
Diphysciales M.Fleisch.
Diphysciaceae M.Fleisch.
Timmiidae Ochyra
Timmiales Ochyra
Timmiaceae Schimp.
Encalyptidae Ochyra, Żarnowiec & Bedn.-Ochyra
Encalyptales Dixon
Encalyptaceae Schimp.
Funariidae Ochyra
Funariales M.Fleisch.
Funariaceae Schwägr.
Disceliaceae Schimp.
Gigaspermidae M.Stech & W.Frey
Gigaspermales Goffinet, Wickett, O.Werner, Ros,
A.J.Shaw & C.J.Cox
Gigaspermaceae Lindb.
Dicranidae Doweld
Catoscopiales Ignatov & Ignatova
Catoscopiaceae Broth.
Dicranales H.Philib ex M.Fleisch.
Timmiellaceae Y.Inoue & H.Tsubota
Distichiaceae Schimp.
Hymenolomataceae Ignatov & Fedosov
Flexitrichaceae Ignatov & Fedosov
Bryoxiphiaceae Besch.
Archidiaceae Schimp.
Micromitriaceae Smyth ex Goffinet &
Budke
Leucobryaceae Schimp.
Amphidiaceae M.Stech
Aongstroemiaceae De Not.
Dicranellaceae M.Stech
Fissidentaceae Schimp.
Dicranaceae Schimp.
Calymperaceae Kindb.
Rhabdoweisiaceae Limpr.
Schistostegaceae Schimp.
Bruchiaceae Schimp.
Ditrichaceae Limpr.
Pottiaceae Schimp.
Grimmiales M.Fleisch.
Saelaniaceae Ignatov & Fedosov
Seligeriaceae Schimp.
Ptychomitriaceae Schimp.
Grimmiaceae Arn.
Bryidae Engl.
Hedwigiales Ochyra
Hedwigiaceae Schimp.
Bartramiales D.Quandt, N.E.Bell & M.Stech
Bartramiaceae Schwägr.
Splachnales Ochyra
Splachnaceae Grev. & Arn.
Meesiaceae Schimp.
8
N. G. HODGETTS ET AL.
Bryales Limpr.
Bryaceae Schwägr.
Mniaceae Schwägr.
Orthotrichales Dixon
Orthotrichaceae Arn.
Orthodontiales N.E.Bell, A.E.Newton & D.Quandt
Orthodontiaceae Goffinet
Aulacomniales N.E.Bell, A.E.Newton & D.Quandt
Aulacomniaceae Schimp.
Rhizogoniales Goffinet & W.R.Buck
Rhizogoniaceae Broth.
Hookeriales M.Fleisch.
Hypopterygiaceae Mitt.
Daltoniaceae Schimp.
Hookeriaceae Schimp.
Leucomiaceae Broth.
Pilotrichaceae Kindb.
Hypnales W.R.Buck & Vitt
Fontinalaceae Schimp.
Plagiotheciaceae M.Fleisch.
Fabroniaceae Schimp.
Pterigynandraceae Schimp.
Habrodontaceae Schimp.
Climaciaceae Kindb.
Myriniaceae Schimp.
Amblystegiaceae G.Roth
Calliergonaceae
Vanderp.,
Hedenäs,
C.J.Cox & A.J.Shaw
Scorpidiaceae Ignatov & Ignatova
Leskeaceae Schimp.
Pseudoleskeaceae Schimp.
Pseudoleskeellaceae Ignatov & Ignatova
Thuidiaceae Schimp.
Brachytheciaceae Schimp.
Hypnaceae Schimp.
Callicladiaceae Jan Kučera & Ignatov
Taxiphyllaceae Ignatov
Pylaisiadelphaceae Goffinet & W.R.Buck
Jocheniaceae Jan Kučera & Ignatov
Stereodontaceae Hedenäs, Schlesak &
D.Quandt
Pylaisiaceae Schimp.
Sematophyllaceae Broth.
Hylocomiaceae M.Fleisch.
Rhytidiaceae Broth.
Entodontaceae Kindb.
Cryphaeaceae Schimp.
Leucodontaceae Schimp.
Antitrichiaceae Ignatov & Ignatova
Neckeraceae Schimp.
Heterocladiellaceae Ignatov & Fedosov
Lembophyllaceae Broth.
Echinodiaceae Broth.
Myuriaceae M.Fleisch.
Anomodontaceae Kindb.
List of Taxa
Subdivisions (‘-phytina’), subclasses (‘-ideae’) and suborders (‘-ineae’), shown in the taxonomic synopsis
above, are not listed in the main checklist, but subfamilies (‘-oideae’) are included. Tribes and subtribes are
included in the Lejeuneaceae, as they are fairly useful
divisions.
Hornworts
ANTHOCEROTOPHYTA
ANTHOCEROTOPSIDA DE BARY EX JANCZ .
Anthocerotales Limpr.
Anthocerotaceae Dumort.
1 Anthoceros L.
1 A. agrestis Paton [Anthoceros punctatus subsp.
agrestis (Paton) Damsh., Anthoceros punctatus var. douinii (R.M.Schust.) Damsh.]
2 A. caucasicus Steph.
3 A. neesii Prosk.
4 A. punctatus L.
Phymatocerotales R.J.Duff
Phymatocerotaceae R.J.Duff
2 Phymatoceros Stotler
1 P. bulbiculosus (Brot.) Stotler, W.T.Doyle &
Crand.-Stotl. [Phaeoceros bulbiculosus
(Brot.) Prosk.]
Notothyladales Hyvönen & Piippo
Notothyladaceae Müll.Frib. ex Prosk.
Notothyloideae Grolle
3 Notothylas Sull.
subgenus Notothylas
1 N. orbicularis (Schwein.) Sull.
Phaeocerotideae Hässel
4 Phaeoceros Prosk.
1 P. carolinianus (Michx.) Prosk. [Phaeoceros
laevis subsp. carolinianus (Michx.)
Prosk.]
2 P. laevis (L.) Prosk.
Liverworts
MARCHANTIOPHYTA
HAPLOMITRIOPSIDA STOTLER & CRAND.- STOTL.
Calobryales Hamlin
Haplomitriaceae Dědeček
1 Haplomitrium Nees
subgenus Haplomitrium
section Haplomitrium
1 H. hookeri (Lyell ex Sm.) Nees
JOURNAL OF BRYOLOGY
a
var. hookeri1
subgenus Sudeticae (Schljakov) L.Söderstr., De Roo &
Hedd. [Pseudolophozia Konstant. & Vilnet]
5 B. sudetica (Nees ex Huebener) L.Söderstr., De
Roo & Hedd. [Lophozia alpestris auct.
non (Schleich. ex F.Weber) A.Evans,
Lophozia sudetica (Nees ex Huebener)
Grolle, Pseudolophozia sudetica (Nees
ex Huebener) Konstant. & Vilnet, Lophozia sudetica var. anomala (Schljakov)
Schljakov,
Lophozia
debiliformis
R.M.Schust. & Damsh., Pseudolophozia
debiliformis (R.M.Schust. & Damsh.) Konstant. & Vilnet, Lophozia debiliformis var.
concolor R.M.Schust. & Damsh.]3
JUNGERMANNIOPSIDA STOTLER & CRAND.- STOTL.
Jungermanniales H.Klinggr.
Adelanthaceae Grolle
Adelanthoideae K.Feldberg
2 Adelanthus Mitt.
section Lindenbergiani Grolle
1 A. lindenbergianus (Lehm.) Mitt.
3
Pseudomarsupidium Herzog
1 P. decipiens (Hook.) Grolle [Adelanthus decipiens
(Hook.) Mitt.]
Jamesonielloideae Inoue
4 Syzygiella Spruce [Jamesoniella (Spruce) F.Lees,
Crossogyna (R.M.Schust.) Schljakov]
subgenus Syzygiella
1 S. autumnalis (DC.) K.Feldberg, Váňa, Hentschel &
Heinrichs [Jamesoniella autumnalis (DC.)
Steph., Crossogyna autumnalis (DC.)
Schljakov]
2 S. rubricaulis (Nees) Steph. [Jamesoniella rubricaulis (Nees) Grolle]
8 Biantheridion (Grolle) Konstant. & Vilnet
1 B. undulifolium (Nees) Konstant. & Vilnet [Jamesoniella undulifolia (Nees) Müll.Frib.]
9 Crossocalyx Meyl
1 C. hellerianus (Nees ex Lindenb.) Meyl. [Anastrophyllum hellerianum (Nees ex
Lindenb.) R.M.Schust.]
10
Gymnocolea (Dumort.) Dumort.
1 G. borealis (Frisvoll & Moen) R.M.Schust.
2 G. fascinifera Potemkin4
3 G. inflata (Huds.) Dumort.
a subsp. acutiloba (Schiffn.) R.M.Schust. ex
L.Söderstr. & Váňa [Gymnocolea
acutiloba (Schiffn.) Müll.Frib.]5
b subsp. inflata [Gymnocolea inflata var. heterostipa (Carringt. ex Spruce) Müll.Frib.]
11
Isopaches H.Buch
1 I. alboviridis (R.M.Schust.) Schljakov6
2 I. bicrenatus (Schmidel ex Hoffm.) H.Buch
[Lophozia bicrenata (Schmidel ex
Hoffm.) Dumort.]
3 I. decolorans (Limpr.) H.Buch [Lophozia decolorans (Limpr.) Steph.]
12
Neoorthocaulis L.Söderstr.
1 N. attenuatus (Mart.) L.Söderstr., De
Hedd. [Barbilophozia attenuata
Loeske, Orthocaulis attenuatus
A.Evans, Lophozia attenuata
Dumort.]
Anastrophyllaceae L.Söderstr., De Roo & Hedd.
5 Anastrepta (Lindb.) Schiffn.
1 A. orcadensis (Hook.) Schiffn.
6
7
1
Anastrophyllum (Spruce) Steph.
1 A. alpinum Steph.2
2 A. assimile (Mitt.) Steph. [Anastrophyllum assimile var. nardioides (Lindb.) Damsh.]
3 A. donnianum (Hook.) Steph.
4 A. joergensenii Schiffn.
5 A. michauxii (F.Weber) H.Buch
Barbilophozia Loeske
subgenus Barbilophozia
1 B. barbata (Schmidel ex Schreb.) Loeske [Lophozia
barbata (Schmidel ex Schreb.) Dumort.]
2 B. hatcheri (A.Evans) Loeske [Lophozia hatcheri
(A.Evans) Steph.]
3 B. lycopodioides (Wallr.) Loeske [Lophozia lycopodioides (Wallr.) Cogn.]
4 B. rubescens (R.M.Schust. & Damsh.) Kartt. &
L.Söderstr.
[Lophozia
rubescens
R.M.Schust. & Damsh.]
9
Roo &
(Mart.)
(Mart.)
Mart.)
Haplomitrium hookeri var. minutum (E.O.Campb.) Barthol.-Began occurs in Japan and New Zealand.
Anastrophyllum alpinum was synonymised with Anastrophyllum joergensenii by Grolle (1964) but reinstated at species level by Long et al. (2006).
3
Lophozia debiliformis was not recognised in Söderström, Hagborg, et al. (2016), following Söderström, De Roo, et al. (2010), although it was sometimes
recognised in later publications. It was nested within Barbilophozia sudetica in the molecular studies by de Roo et al. (2007) and Vilnet, Konstantinova,
et al. (2010) although neither type material nor material from the type locality was included. The ‘taxon’ is common in the mountains of Europe (e.g.
in Scandinavia, the Alps) and elsewhere and may be a modification of harsh environments.
4
Gymnocolea fascinifera was described from Yamal Peninsula and recently recorded for European Russia from Komi Republic by Potemkin (2008).
5
The taxonomic value of Gymnocolea inflata subsp. acutiloba is unclear. A detailed study is needed to clarify whether this is an extreme form of Gymnocolea
inflata s.str.
6
Isopaches alboviridis was recently recorded from European Russia in Leningrad (Potemkin and Rozantseva 2015) and Murmansk Provinces (Borovichev 2008).
2
10
N. G. HODGETTS ET AL.
2
N. binsteadii (Kaal.) L.Söderstr., De Roo &
Hedd. [Barbilophozia binsteadii (Kaal.)
Loeske, Orthocaulis binsteadii (Kaal.)
H.Buch, Lophozia binsteadii (Kaal.)
A.Evans]
3 N. floerkei (F.Weber & D.Mohr) L.Söderstr., De
Roo & Hedd. [Barbilophozia floerkei
(F.Weber & D.Mohr) Loeske, Orthocaulis floerkei (F.Weber & D.Mohr) H.Buch,
Lophozia floerkei (F.Weber & D.Mohr)
Schiffn.]
4 N. hyperboreus (R.M.Schust.) L.Söderstr., De
Roo & Hedd. [Barbilophozia hyperborea
(R.M.Schust.) Potemkin, Orthocaulis
hyperboreus (R.M.Schust.) Konstant.]
13
Orthocaulis H.Buch
1 O. atlanticus (Kaal.) H.Buch [Barbilophozia
atlantica (Kaal.) Müll.Frib., Lophozia
atlantica (Kaal.) Müll.Frib.]
2 O. cavifolius H.Buch & S.W.Arnell [Sphenolobus
cavifolius (H.Buch & S.W.Arnell)
Müll.Frib., Anastrophyllum cavifolium
(H.Buch & S.W.Arnell) Lammes, Lophozia cavifolia (H.Buch & S.W.Arnell)
R.M.Schust.]
14
Schizophyllopsis Váňa & L.Söderstr.
1 S. sphenoloboides (R.M.Schust.) Váňa &
L.Söderstr. [Anastrophyllum sphenoloboides R.M.Schust.]7
15
Schljakovia Konstant. & Vilnet
1 S. kunzeana (Huebener) Konstant. & Vilnet
[Barbilophozia kunzeana (Huebener)
Müll.Frib., Lophozia kunzeana (Huebener) A.Evans]
16
Schljakovianthus Konstant. & Vilnet
1 S. quadrilobus (Lindb.) Konstant. & Vilnet [Barbilophozia quadriloba (Lindb.) Loeske,
Lophozia quadriloba (Lindb.) A.Evans,
Lophozia quadriloba var. glareosa
(Jørg.) Jørg.]
17
Sphenolobopsis R.M.Schust. & N.Kitag.
1 S. pearsonii (Spruce) R.M.Schust.
7
18
Sphenolobus (Lindb.) Berggr.
1 S. minutus (Schreb. ex D.Crantz) Berggr. [Anastrophyllum minutum (Schreb. ex
D.Crantz) R.M.Schust., Anastrophyllum
minutum var. weberi (Mart.) Kartt.]
2 S. saxicola (Schrad.) Steph. [Anastrophyllum
saxicola (Schrad.) R.M.Schust.]
19
Tetralophozia (R.M.Schust.) Schljakov
1 T. filiformis (Steph.) Urmi
2 T. setiformis (Ehrh.) Schljakov
Cephaloziaceae Mig.
Cephalozioideae Müll.Frib.
20 Cephalozia (Dumort.) Dumort.
1 C. ambigua C.Massal. [Cephalozia bicuspidata var.
paludosa Jørg. ex Damsh. nom. inval.]
2 C. bicuspidata (L.) Dumort.
a subsp. bicuspidata
b subsp. lammersiana (Huebener) R.M.Schust.8
3 C. crossii Spruce9
4 C. lacinulata (J.B.Jack ex Gottsche & Rabenh.)
Spruce
5 C. macounii (Austin) Austin
21
Fuscocephaloziopsis Fulford [Pleurocladula Grolle]
1 F. affinis (Lindb. ex Steph.) Váňa & L.Söderstr.
[Cephalozia affinis Lindb. ex Steph.,
Pleurocladula affinis (Lindb. ex Steph.)
Konstant., Vilnet & Troitsky]
2 F. albescens (Hook.) Váňa & L.Söderstr. [Pleurocladula albescens (Hook.) Grolle]
a var. albescens
b var. islandica (Nees) Váňa & L.Söderstr.
[Pleurocladula albescens var. islandica (Nees) L.Söderstr. & Váňa]
3 F. catenulata (Huebener) Váňa & L.Söderstr.
[Cephalozia catenulata (Huebener)
Lindb., Pleurocladula catenulata (Huebener) Konstant., Vilnet & Troitsky]
a subsp. catenulata10
4 F. connivens (Dicks.) Váňa & L.Söderstr. [Cephalozia connivens (Dicks.) Lindb., Pleurocladula
connivens (Dicks.) Konstant., Vilnet &
Trotsky]
a subsp. connivens [Cephalozia connivens
var. compacta (Warnst.) W.E.Nicholson ex Macvicar]11
Schizophyllopsis sphenoloboides was nested within Anastrophyllum in the study by Vilnet, Konstantinova, et al. (2010) which led Stotler and Crandall-Stotler
(2017) to retain it in that genus. As it is the type of Schizophyllopsis, this means that the whole genus would disappear. However, Vilnet, Konstantinova, et al.
(2010) did not include any other member of the genus, or the type of Anastrophyllum, in their studies. We thus retain Schizophyllopsis pending further
studies.
8
Cephalozia bicuspidata subsp. lammersiana has a long history of debate on whether it is worth recognizing at any level. The issue requires a molecular study
before it can be resolved.
9
Cephalozia crossii was reported from the Canary Islands (Schäfer-Verwimp and Váňa in Ellis, Afonina, et al. 2018).
10
Fuscocephaloziopsis catenulata subsp. nipponica (S.Hatt.) Váňa & L.Söderstr. occurs in SE Asia.
11
Fuscocephaloziopsis connivens subsp. fissa (Steph.) Váňa & L.Söderstr. occurs in Africa and SE Asia and subsp. sandwicensis (Mont.) Váňa & L.Söderstr. in
Hawaii, Tahiti and Central America.
JOURNAL OF BRYOLOGY
5
F. crassifolia (Lindenb. & Gottsche) Váňa &
L.Söderstr.
[Cephalozia
crassifolia
(Lindenb. & Gottsche) Fulford]
6 F. leucantha (Spruce) Váňa & L.Söderstr.
[Cephalozia leucantha Spruce, Pleurocladula leucantha (Spruce) Konstant.,
Vilnet & Troitsky, Cephalozia leucantha
var. robusta Schljakov]
7 F. loitlesbergeri (Schiffn.) Váňa & L.Söderstr.
[Cephalozia loitlesbergeri Schiffn., Pleurocladula loitlesbergeri (Schiff.) Konstant., Vilnet & Troitsky]
8 F. lunulifolia (Dumort.) Váňa & L.Söderstr.
[Cephalozia
lunulifolia
(Dumort.)
Dumort., Pleurocladula lunulifolia
(Dumort.) Konstant., Vilnet & Troitsky]
9 F. macrostachya (Kaal.) Váňa & L.Söderstr.
[Cephalozia macrostachya Kaal., Pleurocladula macrostachya (Kaal.) Konstant.,
Vilnet & Troitsky]
a subsp. macrostachya12
i. var. macrostachya
ii. var. spiniflora (Schiffn.) Váňa &
L.Söderstr.
10 F. pleniceps (Austin) Váňa & L.Söderstr. [Cephalozia pleniceps (Austin) Lindb., Pleurocladula pleniceps (Austin) Konstant., Vilnet
& Troitsky]
a var. pleniceps [Cephalozia pleniceps var.
sphagnorum (C.Massal.) Jørg.]13
22
Nowellia Mitt.
section Nowellia
1 N. curvifolia (Dicks.) Mitt.
Odontoschismatoideae H.Buch ex Grolle.
23 Odontoschisma (Dumort.) Dumort. [Cladopodiella
H.Buch]
section Cladopodiella (H.Buch) Gradst., S.C.Aranda
& Vanderp.
1 O. francisci (Hook.) L.Söderstr. & Váňa [Cladopodiella francisci (Hook.) Jørg.]
12
11
section Denudata R.M.Schust.
2 O. denudatum (Mart.) Dumort.
a subsp. denudatum14
3 O. elongatum (Lindb.) A.Evans
4 O. macounii (Austin) Underw.
section Neesii Gradst., S.C.Aranda & Vanderp.
5 O. fluitans (Nees) L.Söderstr. & Váňa [Cladopodiella fluitans (Nees) H.Buch]
section Odontoschisma
6 O. sphagni (Dicks.) Dumort. [Odontoschisma
prostratum (Sw.) Trevis.]15
Cephaloziellaceae Douin16
24 Cephaloziella (Spruce) Schiffn. [Dichiton Mont.]17
subgenus Cephaloziella
1 C. arctogena (R.M.Schust.) Konstant. [Cephaloziella rubella subsp. arctogena
(R.M.Schust.) R.M.Schust. & Damsh.]
2 C. aspericaulis Jørg.
3 C. baumgartneri Schiffn.
4 C. divaricata (Sm.) Schiffn.
a var. divaricata [Cephaloziella divaricata
var.
rupestris
(C.E.O.Jensen)
Damsh. nom. inval.]
b var. scabra (M.Howe) Haynes [Cephaloziella divaricata var. asperifolia
(Taylor) Damsh.]18
5 C. elachista (J.B.Jack ex Gottsche & Rabenh.)
Schiffn.
a var. elachista19
6 C. elegans (Heeg) Schiffn. [Cephaloziella
rubella var. elegans (Heeg) R.M.Schust.]
7 C. grimsulana (J.B.Jack ex Gottsche &
Rabenh.) Lacout.
8 C. hampeana (Nees) Schiffn. ex Loeske
[Cephaloziella hampeana var. subtilis
(Velen) Macvicar]
9 C. massalongi (Spruce) Müll.Frib. [Cephaloziella massalongi var. compacta (Jørg.)
Müll.Frib.]
10 C. nicholsonii Douin
Fuscocephaloziopsis macrostachya subsp. australis (R.M.Schust.) Váňa & L.Söderstr. occurs in SE USA.
Fuscocephaloziopsis pleniceps var. caroliniana (R.M.Schust.) Váňa & L.Söderstr. occurs in SE USA.
14
Odontoschisma denudatum subsp. naviculare (Steph.) Gradst., S.C.Aranda & Vanderp. occurs in E Asia and subsp. sandvicense (Ångstr.) Gradst., S.C.Aranda &
Vanderp. in Japan and Hawaii.
15
Odontoschisma prostratum was recently synonymised by Aranda et al. (2014) based on molecular data.
16
Cephaloziellaceae is here treated in a very broad sense following Váňa et al. (2013). Several genera (Obtusifolium, Oleolophozia, Protolophozia) have recently
been excluded from other families and been indicated to belong closer to Cephaloziellaceae, but there is no comprehensive study of this phylogenetic
region. To avoid monogeneric families of doubtful value the World Checklist (Söderström, Hagborg, et al. 2016) included everything in the only available
family name, Cephaloziellaceae, as a sort of ‘superfamily’, which we are following here.
17
Cephaloziella is one of the most difficult liverwort genera. Almost all taxa are difficult to determine, and their taxonomy and distribution is poorly understood. Bell et al. (2013) provided evidence that the molecular data do not match morphological identifications. Type-based studies using molecular data are
urgently needed.
18
The taxonomic value of Cephaloziella divaricata var. scabra is controversial. In Central Europe var. scabra intergrades with var. divaricata and is possibly only
a morph of shady habitats (Köckinger 2017).
19
Cephaloziella elachista var. spinophylla (C.Gao) C.Gao occurs in China.
13
12
N. G. HODGETTS ET AL.
11
phyllacantha (C.Massal. & Carestia)
Müll.Frib.
12 C. polystratosa (R.M.Schust. & Damsh.) Konstant. [Cephaloziella divaricata var.
polystratosa (R.M.Schust. & Damsh.)
Potemkin]20
13 C. rubella (Nees) Warnst. [Cephaloziella
rubella var. sullivantii (Austin) Müll.Frib.
ex R.M.Schust., Cephaloziella rubella
var. bifida (Lindb.) Douin, Cephaloziella
rubella var. pulchella (C.E.O.Jensen)
R.M.Schust.]
14 C. spinigera (Lindb.) Jørg. [Cephaloziella spinigera f. striatula (C.E.O.Jensen) Damsh.,
Cephaloziella subdentata Warnst.]
15 C. stellulifera (Taylor ex Carrington &
Pearson) Croz. [Cephaloziella stellulifera
var. limprichtii (Warnst.) Macvicar]
16 C. uncinata R.M.Schust.
a var. uncinata21
17 C. varians (Gottsche) Steph. [Cephaloziella
alpina Douin, Cephaloziella arctica
Bryhn & Douin, Cephaloziella varians
var. arctica (Bryhn & Douin) Damsh.,
Cephaloziella varians var. scabra
(S.W.Arnell) Damsh.]
subgenus Dichiton (Mont.) Müll.Frib.
18 C. calyculata (Durieu & Mont.) Müll.Frib.
19 C. integerrima (Lindb.) Warnst. [Dichiton integerrimum (Lindb.) H.Buch, Cephaloziella integerrima var. obtusa Müll.Frib.]
subgenus Evansia (Douin & Schiffn.) Müll.Frib.
20 C. dentata (Raddi) Steph.
subgenus Prionolobus (Spruce) Müll.Frib.
21 C. granatensis (J.B.Jack ex Steph.) Fulford
22 C. turneri (Hook.) Müll.Frib.
25
Obtusifolium S.W.Arnell
1 O. obtusum (Lindb.) S.W.Arnell (Lophozia
obtusa (Lindb.) A.Evans)
26
Oleolophozia L.Söderstr., De Roo & Hedd.
1 O. perssonii (H.Buch & S.W.Arnell) L.Söderstr.,
De Roo & Hedd. [Lophozia perssonii
H.Buch & S.W.Arnell, Lophoziopsis
20
perssonii (H.Buch & S.W.Arnell) Konstant. & Vilnet]
C.
27
Protolophozia (R.M.Schust.) Schljakov
1 P. elongata (Steph.) Schljakov [Lophozia elongata Steph.]
2 P. herzogiana (E.A.Hodgs. & Grolle) Váňa &
L.Söderstr.
[Lophozia
herzogiana
E.A.Hodgs. & Grolle]
Lophoziaceae Cavers
28 Heterogemma (Jørg.) Konstant. & Vilnet [Massularia Schljakov nom. illeg.]
1 H. capitata (Hook.) Konstant. & Vilnet [Lophozia capitata (Hook.) Macoun]
2 H. laxa (Lindb.) Konstant. & Vilnet [Lophozia
laxa (Lindb.) Grolle, Massularia laxa
(Lindb.) Schljakov]
29
Lophozia (Dumort.) Dumort.22
1 L. ascendens (Warnst.) R.M.Schust. [Lophozia
gracillima H.Buch]
2 L. ciliata Damsh., L.Söderstr. & H.Weibull
3 L. fuscovirens Bakalin & Vilnet23
4 L. guttulata (Lindb. & Arnell) A.Evans [Lophozia porphyroleuca (Nees) Schiffn. nom.
illeg., Lophozia longiflora auct. (sensu
Grolle and Long 2000; Söderström,
Urmi, et al. 2002; Damsholt 2002),
Lophozia longiflora var. guttulata
(Lindb. & Arnell) Schljakov]24
5 L. lantratoviae Bakalin25
6 L. longiflora (Nees) Schiffn. [Lophozia ventricosa var. longiflora (Nees) Macoun,
Lophozia ventricosa var. uliginosa
auct. (sensu Söderström, Urmi, et al.
2002; Damsholt 2002)]26
7 L. murmanica Kaal. [Lophozia groenlandica
auct. (sensu Grolle and Long 2000;
Söderström, Urmi, et al. 2002),
Lophozia wenzelii var. groenlandica
auct. (sensu Bakalin 2005; Konstantinova, Bakalin, et al. 2009), Lophozia
confertifolia auct. (sensu Konstantinova, Potemkin, et al. 1992),
Cephaloziella polystratosa was elevated to species rank by Konstantinova (2000b). It is reported from several places in European Russia and Svalbard.
Cephaloziella uncinata var. brevigyna R.M.Schust. & Damsh. and var. sphagnicola R.M.Schust. occur in North America. Schuster (1988; cf. also Damsholt 2013)
also described a var. mamillosa R.M.Schust. & Damsh. from Greenland, but that name is invalid and we do not validate it here.
22
The genus Lophozia includes many taxonomically and nomenclaturally problematic taxa, as outlined in Söderström, Váňa, et al. (2013).
23
Lophozia fuscovirens was recently described from the Russian Far East (Bakalin and Vilnet 2019) and is found also on Svalbard (N. Konstantinova in prep.).
24
For Lophozia guttulata, see footnote on Lophozia longiflora.
25
Lophozia lantratoviae is a recently described species reported from Caucasus by Konstantinova, Akatova, et al. (2009).
26
Since Schljakov (1980), Lophozia longiflora has been used to include Lophozia guttulata, a species mostly restricted to moist dead wood habitats. Lophozia
longiflora was lectotypified by Bakalin (2016). The lectotype corresponds to the concept of Müller (1954), Saukel (1985), Meinunger and Schröder (2007),
Bakalin (2016) and Köckinger (2017), describing a species occurring mostly on peaty soil and in rocky habitats, but not the concept of Grolle and Long
(2000), which corresponds to our Lophozia guttulata.
21
JOURNAL OF BRYOLOGY
8
9
10
11
12
13
14
27
Lophozia heteromorpha R.M.Schust.
& Damsh.]27
L. savicziae Schljakov [Lophozia silvicola var.
grandiretis H.Buch & S.W.Arnell, Lophozia ventricosa var. grandiretis (H.Buch &
S.W.Arnell) R.M.Schust. & Damsh.,
Lophozia murmanica auct. (sensu
Schljakov 1969)]
L. schusteriana Schljakov [Lophozia groenlandica auct. (sensu Schuster 1969)]28
L. silvicola H.Buch [Lophozia ventricosa auct.
(sensu Müller 1954; Schljakov 1980;
Meinunger and Schröder 2007; Köckinger 2017; non Grolle and Long 2000),
Lophozia ventricosa var. silvicola
(H.Buch) E.W.Jones]29
L. silvicoloides N.Kitag.30
L. subapiculata R.M.Schust. & Damsh.31
L. ventricosa (Dicks.) Dumort. [Lophozia
groenlandica auct. (sensu Schljakov
1980, 1998), Lophozia confertifolia
auct. (sensu Schljakov 1975, 1998;
Ştefănuţ 2008), Lophozia murmanica
auct. (sensu Schljakov 1970), Lophozia
ventricosa
var.
confusa
R.M.Schust.]32
L. wenzelii (Nees) Steph. [Lophozia groenlandica auct. (sensu Damsholt 1994,
2002; Ştefănuţ 2008), Lophozia confertifolia Schiffn. (sensu Damsholt 2002;
Köckinger 2017), Lophozia ventricosa
13
var. uliginosa Breidl. ex Schiffn., Lophozia iremelensis Schljakov]33
a var. lapponica H.Buch & S.W.Arnell
b var. litoralis (Arnell) Bakalin
c var. massularioides Bakalin
d var. wenzelii
30
Lophoziopsis Konstant. & Vilnet
1 L. excisa (Dicks.) Konstant. & Vilnet [Lophozia
excisa (Dicks.) Dumort.]
a var. elegans (R.M.Schust.) Konstant. &
Vilnet [Schistochilopsis elegans
(R.M. Schust.) Konstant.]
b var. excisa [Lophozia excisa var. cylindracea
(Dumort.) Müll.Frib.]34
2 L. jurensis (Meyl. ex Müll.Frib.) Mamontov &
Vilnet [Lophozia jurensis Meyl. ex Müll.
Frib., Lophozia propagulifera auct.
eur., Lophoziopsis propagulifera auct.
eur., Lophozia latifolia R.M.Schust.,
Lophoziopsis latifolia (R.M.Schust.)
Köckinger]35
3 L. longidens (Lindb.) Konstant. & Vilnet [Lophozia longidens (Lindb.) Macoun]
a subsp. arctica (R.M.Schust.) Váňa &
L.Söderstr. [Lophoziopsis rubrigemma f. arctica (R.M.Schust.)
Bakalin]
b subsp. longidens
4 L. pellucida (R.M.Schust.) Konstant. & Vilnet
[Lophozia pellucida R.M.Schust.]
Lophozia murmanica was placed in synonymy with Lophozia groenlandica with a question mark (together with Lophozia heteromorpha, also with a question
mark) by Schljakov (1998). This concept has led to questionable reports of Lophozia murmanica from many areas of Europe.
28
Lophozia schusteriana is a new name given to Lophozia groenlandica sensu Schuster (1969) by Schljakov (1975). Schljakov also rejected all reports from
Europe but Vilnet, Konstantinova, et al. (2010) reported it later from Svalbard.
29
For Lophozia silvicola, see footnote under Lophozia ventricosa.
30
Lophozia silvicoloides is originally described from E Asia and recently reported from Svalbard (Vilnet, Milyutina, et al. 2005) and European Russia in Murmansk Province and the Republic of Bashkortostan (Bakalin 2001).
31
Lophozia subapiculata is a critical taxon that was synonymised with Lophozia ventricosa by Bakalin (2005) but recognised as a separate species and reported
from Svalbard by Konstantinova and Savchenko (2018).
32
The Lophozia ventricosa/wenzelii complex has never been studied in detail worldwide using both molecular and morphological methods. Vilnet, Konstantinova, et al. (2010) and de Roo et al. (2007) concentrated on other problems and did not include a sufficient number of specimens of these and related taxa
to include the full morphological variation of this group. The taxonomy proposed here is based mostly on morphological studies (which, however, are
contradictory!) and is therefore provisional pending future research.
The name Lophozia ventricosa has been applied to different concepts since Buch (1929) described Lophozia silvicola (see also Buch 1932) and has often
included the latter as synonym. The neotype chosen by Grolle and Long (2000) for Lophozia ventricosa define it as different from Lophozia silvicola.
However, there are some doubts if this neotype represents Lophozia ventricosa sensu Söderström, Hagborg, et al. (2016) or is a form of Lophozia wenzelii
(see Meinunger and Schröder 2007; Köckinger 2017). Lophozia ventricosa is here understood as a plant with the overall morphology and anatomy of Lophozia silvicola but showing homogeneous granular oil bodies. The neotype of Grolle and Long (2000) needs a reinvestigation as it approaches what we understand as Lophozia wenzelii.
33
Lophozia wenzelii is very close to Lophozia ventricosa according to molecular studies by De Roo et al. (2007) and Vilnet, Konstantinova, et al. (2010), but refer
to the footnote under Lophozia ventricosa concerning the neotype of the latter species. Söderström, Hagborg, et al. (2016) did not accept any subspecific
taxa but we list some here accepted in the revision of Lophozia for Russia (Bakalin 2005) to draw attention to the need for further studies of this complex.
Lophozia groenlandica has been used in at least three different senses (see Söderström, Váňa, et al. 2013). Although being one of the oldest names in
the complex, we refrain from using it in any sense until its affinity is better defined. Schljakov (in Konstantinova, Potemkin, et al. 1992) proposed to reject
the name Lophozia groenlandica (but never formally did it) and changed it for Lophozia confertifolia.
Lophozia confertifolia has also been used in several senses and we here refrain to use it in any specific sense until it is lectotypified (and possibly epitypified if needed).
34
Lophoziopsis excisa var. infuscata (R.M.Schust. & Damsh.) Konstant. & Vilnet and var. succulenta (R.M.Schust. & Damsh.) Konstant. & Vilnet. occur in arctic
North America and Siberia.
35
Lophoziopsis jurensis is a problematic taxon. It was synonymised with Lophozia latifolia by Schljakov (1980) and with Lophozia propagulifera by Bakalin
(2005). Borovichev and Mamontov (2017; see also Köckinger 2017) rejected the synonymy, as European specimens differed molecularily from specimens
named Lophoziopsis propagulifera from Kamchatka. However, there is as yet no study to determine whether Lophoziopsis latifolia (type from arctic North
America), specimens of Lophoziopsis propagulifera from the Southern Hemisphere (type from South Georgia) and Lophoziopsis jurensis (type from the Alps)
are genetically different. We here list only one species, assuming that the Southern Hemisphere taxon is different, and following Schljakov (1980) in regarding Lophoziopsis latifolia as a synonym of Lophoziopsis jurensis.
14
N. G. HODGETTS ET AL.
5
6
31
32
a var. minor (R.M.Schust.) L.Söderstr. & Váňa
b var. pellucida
L. polaris (R.M.Schust.) Konstant. & Vilnet
[Lophozia
polaris
(R.M.Schust.)
R.M.Schust & Damsh.]
a var. polaris
b var. sphagnorum (R.M.Schust.) Konstant. &
Vilnet
L. rubrigemma (R.M.Schust.) Konstant. & Vilnet
[Lophozia rubrigemma R.M.Schust.]
Trilophozia (R.M.Schust.) Bakalin
1 T. quinquedentata (Huds.) Bakalin [Tritomaria
quinquedentata (Huds.) H.Buch, Tritomaria quinquedentata subsp. turgida
(Lindb.) Damsh., Trilophozia quinquedentata subsp. turgida (Lindb.) Konstant., Tritomaria quinquedentata
f. gracilis (C.E.O.Jensen) R.M.Schust.,
Tritomaria
quinquedentata
var.
dentata S.W.Arnell nom. inval., Tritomaria quinquedentata var. grandiretis
H.Buch & S.W.Arnell]
a var. quinquedentata36
Tritomaria Schiffn. ex Loeske
1 T. exsecta (Schmidel) Schiffn. ex Loeske
a subsp. exsecta37
2 T. exsectiformis (Breidl.) Schiffn. ex Loeske
a subsp. arctica R.M.Schust.
b subsp. exsectiformis38
3 T. scitula (Taylor) Jørg.
Scapaniaceae Mig.
33 Diplophyllum (Dumort.) Dumort.
section Diplophyllum
1 D. albicans (L.) Dumort.
section Protodiplophyllum (R.M.Schust.) Váňa &
L.Söderstr.
2 D. obtusatum (R.M.Schust.) R.M.Schust.39
36
3 D.
a
4 D.
a
obtusifolium (Hook.) Dumort.
subsp. obtusifolium40
taxifolium (Wahlenb.) Dumort.
var. taxifolium [Diplophyllum taxifolium
var. macrostictum H.Buch]41
34
Douinia (C.E.O.Jensen) H.Buch
1 D. ovata (Dicks.) H.Buch
35
Pseudotritomaria Konstant. & Vilnet
1 P. heterophylla (R.M.Schust.) Konstant. &
Vilnet42
36
Saccobasis H.Buch
1 S. polita (Nees) H.Buch [Tritomaria polita
(Nees) Jørg.]
2 S. polymorpha (R.M.Schust.) Schljakov [Tritomaria
polita subsp. polymorpha R.M.Schust.]43
37
Scapania (Dumort.) Dumort.
subgenus Gracilidae (H.Buch) Váňa, Hentschel,
Joch.Müll. & Heinrichs
1 S. gracilis Lindb.
subgenus Plicaticalyx Müll.Frib.
section Planifoliae (Müll.Frib.) Potemkin
2 S. nimbosa Taylor
3 S. ornithopodioides44 (With.) Waddell
subgenus Scapania
section Aequilobae (Müll.Frib.) H.Buch
4 S. aequiloba (Schwägr.) Dumort.
5 S. aspera M.Bernet & Bernet
section Apiculatae H.Buch
6 S. apiculata Spruce
7 S. carinthiaca J.B.Jack ex Lindb.45
a var. carinthiaca
b var. massalongi Müll.Frib.
8 S. umbrosa (Schrad.) Dumort.
section Compactae (Müll.Frib.) H.Buch
9 S. compacta (Roth) Dumort.
10 S. kaurinii Ryan
Trilophozia quinquedentata var. assymetrica (Horik.) L.Söderstr. & Váňa occurs in E Asia. Tritomaria quinquedentata is a polymorphic species with many infraspecific taxa described, but the value of them are questioned. The only infrataxon known to be included in a molecular study (subsp. turgida) was nested
among other specimens morphologically belonging to subsp. quinquedentata (De Roo et al. 2007).
37
Tritomaria exsecta subsp. novaezelandia J.J.Engel occurs in New Zealand.
38
Tritomaria exsectiformis subsp. camerunensis S.W.Arnell ex Váňa occurs in Africa.
39
Diplophyllum obtusatum is close to or possibly conspecific with Diplophyllum obtusifolium, differing mainly in being autoicious vs. paroicious. Urmi (2017)
regarded them as conspecific but did not formally treat them as such. In addition, Bakalin and Vilnet (2018) treated all reports of both species from Asia as
belonging to their new species, Diplophyllum sibiricum Vilnet & Bakalin, and regarded Diplophyllum obtusatum as an American taxon of unclear status.
However, they did not include any American or European specimen of Diplophyllum obtusatum to confirm this conclusion.
40
Diplophyllum obtusifolium subsp. domesticum (Gottsche) Váňa occurs in the Subantarctic.
41
Diplophyllum taxifolium var. mucronatum R.M.Schust. occurs in North America.
42
Pseudotritomaria heterophylla is reported from Franz Josef Land but “the material is poor” (Konstantinova and Potemkin 1996) and the taxon was not
included in the list of Grolle and Long (2000). We include it here with some hesitation.
43
Saccobasis polymorpha was included (as Tritomaria polymorpha) in Tritomaria polita by Grolle and Long (2000).
44
Withering (1776) originally spelt the species epithet of Scapania ornithopodioides as “ornithopoides”, a spelling Potemkin (2019) argued should be retained,
referring to ICN Art. 60.1 (Turland et al. 2018). However, Grolle (1973) argued that “ornithopoides” is bad Latin and in addition that Withering referred to
Dillenius’ pre-Linnean name “… ornithopodii minor …”. Thus Grolle (1973) rejected the original spelling as an orthographic or typographic error, which has
been widely accepted and is followed here.
45
There are two morphotypes of Scapania carianthiaca (Scapania carinthiaca s.str. and Scapania massalongi), which were for a long time treated as separate
taxa (Müller 1954), before being synonymised by Potemkin (1999). Most records in Europe belong to the massalongi-morphototype, while the carinthiacamorphotype is very rare. Further studies with fertile plants are desirable, especially as two morphotypes can be distinguished in the Alps, following Meinunger and Schröder (2007) and others.
JOURNAL OF BRYOLOGY
11 S. spitsbergensis (Lindb.) Müll.Frib.
section Curtae (Müll.Frib.) H.Buch
12 S. curta (Mart.) Dumort.
a var. curta
b var. grandiretis R.M.Schust.
c var. isoloba R.M.Schust.
13 S. helvetica Gottsche
14 S. irrigua (Nees) Nees
a subsp. irrigua [Scapania irrigua var. rubescens H.Buch]
b subsp. rufescens (Loeske) R.M.Schust.
15 S. lingulata H.Buch
a var. lingulata
b var. microphylla (Warnst.) R.M.Schust.
16 S. mucronata H.Buch
17 S. obcordata (Berggr.) S.W.Arnell [Scapania
paradoxa R.M.Schust.]
18 S. parvifolia Warnst.
19 S. praetervisa Meyl. [Scapania mucronata
subsp. praetervisa (Meyl.) R.M.Schust.]
20 S. scandica (Arnell & H.Buch) Macvicar
a var. argutedentata H.Buch
b var. grandiretis (Schljakov) Schljakov
c var. scandica46
21 S. uliginosa (Lindenb.) Dumort.
22 S. zemliae S.W.Arnell
section Cuspiduligerae H.Buch
23 S. cuspiduligera (Nees) Müll.Frib.
a var. cuspiduligera47
section Hyperboreae Váňa, Hentschel, Joch.Müll. &
Heinrichs
24 S. hyperborea Jørg.
25 S. paludicola Loeske & Müll.Frib. [Scapania
paludicola var. rufescens Damsh. nom.
inval.]
a var. paludicola48
b var. rotundiloba R.M.Schust. ex Konstant. &
L.Söderstr.
26 S. tundrae (Arnell) H.Buch
section Kaalaasia (H.Buch) Jørg.
27 S. calcicola (Arnell & J.Perss.) Ingham
28 S. gymnostomophila Kaal.
29 S. ligulifolia R.M.Schust.
section Nemorosae (Müll.Frib.) H.Buch
30 S. crassiretis Bryhn
31 S. degenii Schiffn. ex Müll.Frib. [Scapania brevicaulis auct. eur. non Taylor, Scapania
degenii var. dubia R.M.Schust., Scapania brevicaulis var. dubia (R.M.Schust.)
Damsh.]49
46
15
32 S. nemorea (L.) Grolle
section Scapania
33 S. obscura (Arnell & C.E.O.Jensen) Schiffn.
34 S. paludosa (Müll.Frib.) Müll.Frib. [Scapania
paludosa var. isoloba Müll.Frib., Scapania paludosa var. rubiginosa Müll.Frib.,
Scapania paludosa var. vogesiaca
Müll.Frib.]
35 S. subalpina (Nees ex Lindenb.) Dumort.
a var. subalpina50
36 S. undulata (L.) Dumort. [Scapania undulata
var. aequatiformis (De Not.) C.Massal.
& Carestia, Scapania undulata var.
dentata (Dumort.) Jørg., Scapania
undulata var. oakesii (Austin) H.Buch]
section Scapaniella (H.Buch) Potemkin
37 S. glaucocephala (Taylor) Austin
a var. glaucocephala51
38 S. scapanioides (C.Massal.) Grolle [Scapania
glaucocephala
var.
scapanioides
(C.Massal.) Damsh.]
section Simmonsiae (R.M.Schust.) Váňa, Hentschel,
Joch.Müll. & Heinrichs
39 S. simmonsii Bryhn & Kaal.
section Sphaeriferae Konstant. & Potemkin
40 S. sphaerifera H.Buch & Tuom.
section Verrucosae Potemkin
41 S. verrucosa Heeg
38
Schistochilopsis (N.Kitag.) Konstant.
1 S. grandiretis (Lindb. ex Kaal.) Konstant.
[Lophozia grandiretis (Lindb. ex
Kaal.) Schiffn., Lophozia grandiretis
var. parviretis R.M.Schust., Lophozia
grandiretis var. proteidea (Arnell)
Arnell]
2 S. hyperarctica Konstant. & L.Söderstr. [Lophozia hyperarctica R.M.Schust.]
3 S. incisa (Schrad.) Konstant. [Lophozia incisa
(Schrad.) Dumort., Massularia incisa
(Schrad.) Schljakov, Schistochilopsis
incisa var. inermis (Müll.Frib.) Konstant.]
4 S. opacifolia (Culm. ex Meyl.) Konstant.
[Lophozia opacifolia Culm. ex Meyl.,
Lophozia incisa subsp. opacifolia
(Culm. ex Meyl.) R.M.Schust. & Damsh.]
Acrobolbaceae E.A.Hodgs.
Acrobolboideae R.M.Schust. ex Briscoe
Scapania scandica var. dimorpha R.M.Schust. occurs in North America
Scapania cuspiduligera var. diplophyllopsis R.M.Schust. occurs in North America.
48
Scapania paludicola var. viridigemma R.M.Schust. occurs in North America.
49
Scapania degenii was synonymised with Scapania brevicaulis by Potemkin (1998) but recognised by Konstantinova, Bakalin, et al. (2009). Scapania brevicaulis s.str. is an American taxon that has never been reported from Europe without Scapania degenii being included as as a synonym.
50
Scapania subalpina var. haynesiae Frye & L.Clark and var. muddiae C.D.Bird & W.S.Hong occur in North America.
51
Scapania glaucocephala var. saxicola (R.M.Schust.) Potemkin occurs in North America.
47
16
39
N. G. HODGETTS ET AL.
Acrobolbus Nees [Tylimanthus Mitt.]
1 A. azoricus (Grolle & Perss.) Briscoe [Tylimanthus azoricus Grolle & Perss.]
2 A. madeirensis (Grolle & Perss.) Briscoe [Tylimanthus madeirensis Grolle & Perss.]
3 A. wilsonii Nees
a var. wilsonii52
Antheliaceae R.M.Schust.
40 Anthelia (Dumort.) Dumort.
1 A. julacea (L.) Dumort.
2 A. juratzkana (Limpr.) Trevis.
Arnelliaceae Nakai
41 Arnellia Lindb.
1 A. fennica (Gottsche) Lindb.
Calypogeiaceae Arnell
42 Calypogeia Raddi
subgenus Asperifoliae (Warnst.) R.M.Schust.
1 C. arguta Nees & Mont.
subgenus Calypogeia
2 C. azorica Bischl.
3 C. azurea Stotler & Crotz
4 C. fissa (L.) Raddi
a subsp. fissa
b subsp. neogaea R.M.Schust.
c var. paludosa (Warnst.) Damsh.53
5 C. integristipula Steph.
6 C. muelleriana (Schiffn.) Müll.Frib.
a subsp. muelleriana [Calypogeia muelleriana
var. erecta (Müll.Frib.) Müll.Frib.]54
7 C. neesiana (C.Massal. & Carestia) Müll.Frib.
a subsp. neesiana [Calypogeia neesiana var.
hygrophila Müll.Frib., Calypogeia
neesiana var. repanda (Müll.Frib.)
Meyl.]55
8 C. sphagnicola (Arnell & J.Perss.) Warnst. &
Loeske
9 C. suecica (Arnell & J.Perss.) Müll.Frib.
43
52
Mnioloma Herzog
subgenus Caracoma (Bischl.) R.M.Schust.
1 M. fuscum (Lehm.) R.M.Schust.
Endogemmaceae Konstant.56
44 Endogemma Konstant.
1 E. caespiticia (Lindenb.) Konstant., Vilnet &
A.V.Troitsky [Jungermannia caespiticia
Lindenb.]
Geocalycaceae H.Klinggr.
45 Geocalyx Nees
1 G. graveolens (Schrad.) Nees
Gymnomitriaceae H.Klinggr.
Gymnomitrioideae T.Jensen
46 Gymnomitrion Corda [Apomarsupella R.M.Schust.]
1 G. adustum Nees [Marsupella adusta (Nees)
Spruce]
2 G. alpinum (Gottsche ex Husn.) Schiffn. [Marsupella alpina (Gottsche ex Husn.)
Bernet]
3 G. brevissimum (Dumort.) Warnst. [Marsupella
brevissima (Dumort.) Grolle]
4 G. commutatum (Limpr.) Schiffn. [Marsupella
commutata (Limpr.) Bernet]
5 G. concinnatum (Lightf.) Corda
6 G. corallioides Nees
7 G. crenulatum Gottsche ex Carrington
8 G. obtusum Lindb.
9 G. revolutum (Nees) H.Philib. [Apomarsupella
revoluta (Nees) R.M.Schust.]
a subsp. revolutum57
47
Marsupella Dumort.
1 M. andreaeoides (Lindb.) Müll.Frib.
2 M. apiculata Schiffn. [Gymnomitrion apiculatum (Schiffn.) Müll.Frib.]
3 M. aquatica (Lindenb.) Schiffn. [Marsupella
emarginata subsp. aquatica (Lindenb.)
Meyl., Marsupella emarginata var.
aquatica (Lindenb.) Dumort.]58
4 M. arctica (Berggr.) Bryhn & Kaal.
5 M. boeckii (Austin) Lindb. ex Kaal.
6 M. condensata (Ångstr. ex C.Hartm.) Lindb.
ex Kaal.
7 M. emarginata (Ehrh.) Dumort. [Marsupella
emarginata var. pearsonii (Schiffn. ex
Macvicar) Jørg.]
Acrobolbus wilsonii var. andinus Spruce occurs in South America. Whether it belongs to Acrobolbus wilsonii or some other species is unclear, as Acrobolbus
wilsonii is normally considered to be a European endemic.
Damsholt (2017) transferred Calypogeia sphagnicola f. paludosa (Warnst.) R.M.Schust. to Calypogeia fissa. Buczkowska et al. (2018) included both ‘f. sphagnicola’ and ‘f. paludosa’ in their study on, primarily, the Calypogeia species with blue oil bodies. However, in their molecular phylogenetic tree, ‘paludosa’
came out distinctly separated from ‘sphagnicola’ and closer to Calypogeia fissa (supporting Damsholt’s view), although far enough from it that it should
perhaps deserve recognition at species level. For now, to avoid premature nomenclatural changes we treat it as a variety without affiliation to any of the
subspecies pending further research.
54
Calypogeia muelleriana subsp. blomquistii R.M.Schust. occurs in North America.
55
Calypogeia neesiana subsp. subalpina (Inoue) Inoue occurs in Japan.
56
Endogemma is feminine not neuter and the family name should therefore be Endogemmaceae, not Endogemmataceae as originally published.
57
Gymnomitrion revolutum subsp. novoguineanense (R.M.Schust.) Váňa, Crand.-Stotl. & Stotler occurs in New Guinea.
58
Marsupella aquatica was treated under Marsupella emarginata in Grolle and Long (2000) but shown to be sister to it and accepted at species level by Vilnet,
Konstantinova, et al. (2010).
53
JOURNAL OF BRYOLOGY
8 M. funckii (F.Weber & D.Mohr) Dumort.
[Marsupella ramosa Müll.Frib., Marsupella funckii var. badensis (Schiffn.)
Fam.]
9 M. profunda Lindb.
10 M. sparsifolia (Lindb.) Dumort.
a subsp. sparsifolia59
11 M. sphacelata (Giesecke ex Lindenb.)
Dumort.
12 M. spiniloba R.M.Schust. & Damsh.
13 M. sprucei (Limpr.) Bernet [Marsupella sprucei
var. neglecta (Limpr.) Damsh., Marsupella sprucei var. ustulata (Limpr.)
Damsh.]
14 M. stableri Spruce
15 M. subemarginata Bakalin & Fedosov60
16 M. tubulosa Steph. [Marsupella emarginata subsp. tubulosa (Steph.) N.Kitag.] 61
48
Prasanthus Lindb.
1 P. suecicus (Gottsche) Lindb.
Nardioideae Váňa
49 Nardia Gray
1 N. breidleri (Limpr.) Lindb.
2 N. compressa (Hook.) Gray
3 N. geoscyphus (De Not.) Lindb.
a var. bifida R.M.Schust.62
b var. geoscyphus63
c var. suberecta (Lindb. ex Kaal.) Váňa
4 N. insecta Lindb.
5 N. japonica Steph.
6 N. pacifica Bakalin64
7 N. scalaris Gray
a var. scalaris65
Harpanthaceae Arnell
50 Harpanthus Nees
1 H. flotovianus (Nees) Nees [Harpanthus flotovianus var. chiloscyphoides C.E.O.Jensen, Harpanthus flotovianus var.
latifolia Jørg., Harpanthus flotovianus
var. retusa Jørg.]
59
17
2 H. scutatus (F.Weber & D.Mohr) Spruce
Hygrobiellaceae Konstant. & Vilnet
51 Hygrobiella Spruce
1 H. laxifolia (Hook.) Spruce
Jungermanniaceae Rchb.
Delavayelloideae Grolle
52 Liochlaena Nees
1 L. lanceolata Nees [Jungermannia lanceolata
auct. non L., Jungermannia leiantha
Grolle, Jungermannia subulata var.
leiantha (Grolle) Damsh.]
2 L. subulata (A.Evans) Schljakov [Jungermannia
subulata A.Evans]
Jungermannioideae Dumort.
53 Eremonotus Lindb. & Kaal. ex Pearson
1 E. myriocarpus (Carrington) Lindb. & Kaal. ex
Pearson
54
Jungermannia L.
1 J. atrovirens Dumort. [Jungermannia lanceolata L. nom. rejic., Jungermannia lanceolata var. atrovirens (Dumort.)
Damsh.]
2 J. borealis Damsh. & Váňa
3 J. calcicola Konstant. & Vilnet66
4 J. eucordifolia Schljakov [Jungermannia exsertifolia subsp. cordifolia (Dumort.) Váňa]67
5 J. polaris Lindb. [Jungermannia pumila subsp.
polaris (Lindb.) R.M.Schust.]
6 J. pumila With. [Jungermannia pumila var.
alpestris Gottsche & Rabenh.]
Mesoptychioideae R.M.Schust.
55 Mesoptychia (Lindb.) A.Evans [Leiocolea (Müll.Frib.) Buch]
1 M. badensis (Gottsche ex Rabenh.) L.Söderstr. &
Váňa [Lophozia badensis (Gottsche ex
Rabenh.) Schiffn., Leiocolea badensis
(Gottsche ex Rabenh.) Jørg., Lophozia
badensis var. obtusiloba (Bernet) Schiffn.]
Marsupella sparsifolia subsp. childii R.M.Schust. occurs in New Zealand.
Marsupella subemarginata was described by Bakalin et al. (2019) from Kamchatka but also reported from Switzerland.
61
Marsupella tubulosa was treated as a subspecies of Marsupella emarginata by Söderström, Hagborg, et al. (2016) but shown to be better treated at species
level by Bakalin et al. (2019).
62
Nardia geoscyphus var. bifida was not recognised by Söderström, Hagborg, et al. (2016) but we list it here pending better understanding of the variability of
the species.
63
Nardia geoscyphus var. dioica Bakalin occurs in E Asia.
64
Nardia pacifica is a newly described species (Bakalin and Klimova 2016).
65
Nardia scalaris var. botryoidea (R.M.Schust.) Váňa occurs in North America and var. harae (Amakawa) Váňa in E Asia.
66
Jungermannia calcicola is a newly described species from Russian Caucasus (Konstantinova and Vilnet 2016) and recently also recorded from Albania by
Marka et al. (2018).
67
Jungermannia eucordifolia was treated as a subspecies (Jungermannia exsertifolia subsp. cordifolia) in Söderström, Hagborg, et al. (2016). However, Konstantinova and Vilnet (2016) and later Mamontov, Konstantinova, et al. (2018) showed that it deserves specific status, as was proposed by Schljakov (1981)
and followed by Russian bryologists (e.g. Konstantinova, Potemkin, et al. 1992; Konstantinova, Bakalin, et al. 2009).
60
18
N. G. HODGETTS ET AL.
2
M. bantriensis (Hook.) L.Söderstr. & Váňa
[Lophozia bantriensis (Hook.) Steph.,
Leiocolea bantriensis (Hook.) Jørg.]
a subsp. bantriensis [Lophozia bantriensis
var. subcompressa (Limpr.) Damsh.
ex L.Söderstr.]
b subsp. wallfischii (Ştefănuţ) L.Söderstr. &
Váňa [Leiocolea bantriensis subsp.
wallfischii Ştefănuţ]68
3 M. collaris (Nees) L.Söderstr. & Váňa [Leiocolea
collaris (Nees) Schljakov, Lophozia
alpestris (Schleich ex F.Weber) Evans
nom. rej., Leiocolea alpestris (Schleich.
ex F.Weber) Isov., Lophozia alpestris
var. libertae (Huebener) Damsh.]
4 M. fitzgeraldiae (Paton & A.R.Perry) L.Söderstr. &
Váňa [Leiocolea fitzgeraldiae Paton &
A.R.Perry]
5 M. gillmanii (Austin) L.Söderstr. & Váňa [Lophozia
gillmanii (Austin) R.M.Schust., Leiocolea
gillmanii (Austin) A.Evans, Lophozia gillmanii var. acutifolia (Limpr.) R.M.Schust.]
a var. gillmanii
b var. laxa (Schiffn. ex Burrell) L.Söderstr. [Leiocolea rutheana var. laxa (Schiffn. ex
Burrell) Paton, Lophozia rutheana
var. laxa (Schiffn. ex Burrell) Paton
ex Damsh. nom. inval.]69
6 M. heterocolpos (Thed. ex Hartm.) L.Söderstr.
& Váňa [Lophozia heterocolpos (Thed.
ex Hartm.) M.Howe, Leiocolea heterocolpos (Thed. ex Hartm.) H.Buch]
a var. arctica (S.W.Arnell) L.Söderstr. &
Váňa [Lophozia heterocolpos var.
arctica (S.W.Arnell) R.M.Schust. &
Damsh.]
b var. harpanthoides (Bryhn & Kaal.)
L.Söderstr. & Váňa [Lophozia heterocolpos
var.
harpanthoides
(Bryhn & Kaal.) R.M.Schust.]
c var. heterocolpos
7 M. rutheana (Limpr.) L.Söderstr. & Váňa
[Lophozia ruthana (Limpr.) M.Howe,
Leiocolea rutheana (Limpr.) Müll.Frib.]
8 M. sahlbergii (Lindb. & Arnell) A.Evans
9 M. turbinata (Raddi) L.Söderstr. & Váňa [Leiocolea turbinata (Raddi) H.Buch]
68
Saccogynaceae Heeg
56 Saccogyna Dumort.
1 S. viticulosa (L.) Dumort.
Solenostomataceae Stotler & Crand.-Stotl.
57 Cryptocolea R.M.Schust.
1 C. imbricata R.M.Schust.
58
Solenostoma Mitt. [Plectocolea (Mitt.) Mitt.]
subgenus Eucalyx (Lindb.) Váňa, Crand.-Stotl. &
Stotler
1 S. obovatum (Nees) C.Massal. [Jungermannia
obovata Nees, Plectocolea obovata
(Nees) Mitt.]
2 S. subellipticum (Lindb. ex Heeg) R.M.Schust.
[Jungermannia subelliptica (Lindb. ex
Heeg) Levier, Plectocolea subelliptica
(Lindb. ex Heeg) A.Evans, Jungermannia
obovata subsp. minor (Carrington)
Damsh.]70
subgenus Metasolenostoma Váňa, Crand.-Stotl. &
Stotler
3 S. gracillimum (Sm.) R.M.Schust. [Jungermannia gracillima Sm., Jungermannia
gracillima var. crenulata (Mitt.) Damsh.]
4 S. handelii (Schiffn.) Müll.Frib. [Jungermannia
handelii (Schiffn.) Amakawa]
subgenus Plectocolea Mitt.
5 S. callithrix (Lindenb. & Gottsche) Steph.
[Jungermannia callithrix Lindenb. &
Gottsche]
6 S. hyalinum (Lyell) Mitt. [Jungermannia hyalina
Lyell, Plectocolea hyalina (Lyell) Mitt.]
7 S. paroicum (Schiffn.) R.M.Schust. [Jungermannia paroica (Schiffn.) Grolle]
subgenus Solenostoma
8 S. caucasicum (Váňa) Konstant.71
9 S. confertissimum (Nees) Schljakov [Jungermannia confertissima Nees]
10 S. sphaerocarpum (Hook.) Steph. [Jungermannia sphaerocarpa Hook., Solenostoma
pusillum (C.E.O.Jens.) Steph., Jungermannia jenseniana Grolle, Jungermannia
sphaerocarpa var. nana (Nees ex Flot.)
Frye & L.Clark nom. illeg., Solenostoma
sphaerocarpum var. nanum (Nees ex
Flot.) R.M.Schust.]
Mesoptychia bantriensis subsp. wallfischii was described as Leiocolea bantriensis subsp. wallfischii by Ştefănuţ (2008) from Romania.
Mesoptychia gillmanii var. laxa was accepted by Söderström, Hagborg, et al. (2016) as Mesoptychia rutheana var. laxa although with low confidence.
However, the study by Bell et al. (2013) was largely overlooked and the relation to Mesoptychia gillmanii rather than to Mesoptychia rutheana was not
acknowledged.
70
Solenostoma subellipticum was nested in Solenostoma obovatum in Shaw et al. (2015) and thus reduced to a synonym, a treatment followed by Söderström,
Hagborg, et al. (2016). However, as it seems distinct in at least parts of its distribution area we list it here with some hesitation.
71
Solenostoma caucasicum was reported as new to Europe from the northern part of Caucasus by Konstantinova and Savchenko (2013).
69
JOURNAL OF BRYOLOGY
Southbyaceae Váňa, Crand.-Stotl., Stotler & D.G.Long
59 Gongylanthus Nees
1 G. ericetorum (Raddi) Nees
60
Southbya Spruce
1 S. nigrella (De Not.) Henriq.
2 S. tophacea (Spruce) Spruce
Blepharostomataceae W.Frey & M.Stech
61 Blepharostoma (Dumort.) Dumort.
1 B. trichophyllum (L.) Dumort.
a subsp. brevirete (Bryhn & Kaal.) R.M.Schust.
[Blepharostoma trichophyllum var.
brevirete Bryhn & Kaal.]
b subsp. trichophyllum
Herbertaceae Müll.Frib. ex Fulford & Hatcher
62 Herbertus Gray
1 H. azoricus (Steph.) P.W.Richards72
2 H. borealis Crundw. [Herbertus delavayi auct.
eur. non (Steph.) Steph.]
3 H. hutchinsiae (Gottsche & Rabenh.) A.Evans
[Herbertus aduncus auct. non (Dicks.)
Gray, Herbertus aduncus subsp. hutchinsiae
(Gottsche
&
Rabenh.)
R.M.Schust. nom. inval.]73
4 H. norenus D.G.Long, D.Bell & H.H.Blom74
5 H. sendtneri (Nees) Lindb.
6 H. stramineus (Dumort.) Trevis.
Lepidoziaceae Limpr.
Bazzanioideae Rodway
63 Bazzania Gray
1 B. azorica H.Buch & Perss.
2 B. flaccida (Dumort.) Grolle
3 B. pearsonii Steph.
4 B. tricrenata (Wahlenb.) Lindb.
a var. tricrenata75
72
19
5 B. trilobata (L.) Gray
a var. depauperata (Müll.Frib.) Grolle
b var. trilobata
Lembidioideae R.M.Schust.
64 Kurzia G.Martens
1 K. pauciflora (Dicks.) Grolle
2 K. sylvatica (A.Evans) Grolle
3 K. trichoclados (Müll.Frib.) Grolle
Lepidozioideae Müll.Frib.
65 Lepidozia (Dumort.) Dumort.
1 L. cupressina (Sw.) Lindenb.
a subsp. cupressina [Lepidozia cupressina
subsp. pinnata (Hook.) Pócs, Lepidozia cupressina var. dissitifolia
(Jørg.) Damsh.]76
2 L. pearsonii Spruce
3 L. reptans (L.) Dumort. [Lepidozia reptans var.
julacea (Nees) Damsh. nom. inval., Lepidozia reptans var. tenera (Huebener)
P.Allorge nom. inval.]
4 L. stuhlmannii Steph.77
66
Tricholepidozia (R.M.Schust.) E.D.Cooper
1 T. lindenbergii (Gottsche) E.D.Cooper78
a var. lindenbergii79
2 T. tetradactyla (Hook.f. & Taylor) E.D.Cooper
[Telaranea longii Paton, Telaranea murphyae Paton]80
Zoopsidoideae R.M.Schust.
67 Telaranea Spruce ex Schiffn.
1 T. azorica (H.Buch & Perss.) Pócs [Lepidozia
azorica H.Buch & Perss.]
2 T. europaea J.J.Engel & G.L.Merr. [Telaranea
nematodes auct. eur. non (Gottsche
ex Austin) M.Howe]
Only a single species of Herbertus, Herbertus azoricus, has to date been published from the Azores. However, Richards (1936) observed that there are two
’forms’ present in the archipelago, and it is now clear that two species are present (Hodgetts and Rumsey in prep.), but further analysis is needed to characterize them. Feldberg et al. (2004) suggested that Herbertus azoricus should be placed in synonymy with Herbertus sendtneri. However, further studies show
that spurious lumping of taxa in Herbertus gives a much too simplified view of this complex genus (F. Rumsey pers. comm. 2018), and Herbertus sendtneri is
therefore best regarded as an exclusively Northern Hemisphere species, restricted to the Austrian Alps, Himalaya, and the Pacific Northwest of North
America. Current indications are that Herbertus azoricus is best treated as a synonym or a subspecies of Herbertus juniperoideus, that Herbertus sendtneri
does not occur in Macaronesia, and that this second Azorean species has affinities with South American taxa and plants from the Appalachians previously
assigned to Herbertus tenuis.
73
Reports of Herbertus aduncus from Europe belong to Herbertus hutchinsiae, treated for a long time as a synonym or subspecies of Herbertus aduncus.
74
Herbertus norenus was described from Norway and Scotland by Bell et al. (2012).
75
Bazzania tricrenata var. fulfordiae W.S.Hong occurs in North America.
76
Lepidozia cupressina subsp. africana (Steph.) Pócs occurs in Africa and subsp. natalensis (Steph.) Pócs in southern Africa.
77
Lepidozia stuhlmannii was reported new to the Azores by Frahm (2005). Lepidozia stuhlmannii subsp. pulvinata (Steph.) Pócs and var. carnosa (Steph.) Pócs
occur in Africa. The subspecific identity of the plants in the Azores is unknown.
78
Tricholepidozia lindenbergii was reported as new to Britain (and the Northern Hemisphere) by Blackstock et al. (2019).
79
Trichlepidozia lindenbergii var. complanata (J.J.Engel & G.L.Merr.) E.D.Cooper, var. mellea (J.J.Engel & G.L.Merr.) E.D.Cooper and var. papillata (J.J.Engel &
G.L.Merr.) E.D.Cooper, as well as var. lindenbergii occur in New Zealand.
80
Tricholepidozia was elevated to a separate genus by Cooper et al. (2013) and Tricholepidozia tetradactyla was shown to be the same as Telaranea longii
(Engel and Smith Merrill 2004) and Telaranea murphyae (Villarreal et al. 2014) from Britain.
20
N. G. HODGETTS ET AL.
a subsp. fragrans86
6 L. heterophylla (Schrad.) Dumort. [Chiloscyphus profundus (Nees) J.J.Engel &
R.M.Schust.]
a subsp. heterophylla87
7 L. minor Nees [Chiloscyphus minor (Nees)
J.J.Engel & R.M.Schust.]
8 L. semiteres (Lehm.) Mitt.
a var. semiteres88
Lophocoleaceae Vanden Berghen
68 Chiloscyphus Corda81
1 C. pallescens (Ehrh.) Dumort.
a var. fragilis (Roth) Müll.Frib. [Chiloscyphus
fragilis (Roth) Schiffn.]
b var. pallescens
2 C. polyanthos (L.) Corda
a var. polyanthos
b var. rivularis (Schrad.) Lindb. & Arnell
[Chiloscyphus rivularis (Schrad.) Hazl.]
69
Heteroscyphus Schiffn.
1 H. denticulatus (Mitt.) Schiffn.
2 H. fissistipus (Hook.f. & Taylor) Schiffn.82
Mastigophoraceae R.M.Schust.
72 Mastigophora Nees
1 M. woodsii (Hook.) Nees
70
Leptoscyphus Mitt.
subgenus Anomylia (R.M.Schust.) R.M.Schust.
1 L. cuneifolius (Hook.) Mitt.
subgenus Leptoscyphus
section Leptoscyphus
2 L. porphyrius (Nees) Grolle
a subsp. azoricus (H.Buch & Perss.) Vanderp.
& Heinrichs [Leptoscyphus azoricus
(H.Buch & Perss.) Grolle]83
Plagiochilaceae Müll.Frib.
73 Pedinophyllum Lindb. ex Nordst.
1 P. interruptum (Nees) Kaal.
71
81
Lophocolea (Dumort.) Dumort.
1 L. bidentata (L.) Dumort. [Lophocolea cuspidata (Nees) Limpr.]84
2 L. bispinosa (Hook.f. & Taylor) Gottsche, Lindenb.
& Nees
3 L. brookwoodiana Paton & Sheahan85
4 L. coadunata (Sw.) Mont. [Chiloscyphus coadunatus (Sw.) J.J.Engel & R.M.Schust.,
Lophocolea cuspidata auct. non (Nees)
Limpr.], Lophocolea bidentata var. rivularis (Raddi) Schiffn., Chiloscyphus latifolius (Nees) J.J.Engel & R.M.Schust.]
5 L. fragrans (Moris & De Not.) Gottsche, Lindenb.
& Nees [Chiloscyphus fragrans (Moris &
De Not.) J.J.Engel & R.M.Schust.]
74
Plagiochila (Dumort.) Dumort.
section Arrectae Carl
1 P. bifaria (Sw.) Lindenb.
a var. bifaria89
2 P. papillifolia Steph.90
3 P. punctata (Taylor) Taylor
4 P. retrorsa Gottsche
5 P. spinulosa (Dicks.) Dumort.
6 P. stricta Lindenb.
section Fuscoluteae Carl
7 P. heterophylla Lindenb.
a var. heterophylla [Plagiochila atlantica
F.Rose]91
section Glaucescentes Carl
8 P. longispina Lindenb. & Gottsche
section Plagiochila
9 P. arctica Bryhn & Kaal. [Plagiochila asplenioides subsp. arctica (Bryhn & Kaal.)
R.M.Schust.]
a var. arctica [?Plagiochila asplenioides var.
lobata (Kaal.) Jørg.]92
The Chiloscyphus polyanthos complex has been treated in various ways in the past, without any recent solution. Some authors have treated it as one species
(Chiloscyphus polyanthos) with two subspecies (subsp. polyanthos and pallescens (e.g. Smith 1990); others as four segregate species (Chiloscyphus polyanthos,
Chiloscyphus pallescens, Chiloscyphus fragilis and Chiloscyphus rivularis (e.g. Konstantinova, Potemkin, et al. 1992). Grolle and Long (2000) kept two species, but
did not deal with any subspecific taxa. Söderström, Urmi, et al. (2002) kept two species but recognised four varieties (Chiloscyphus polyanthos var. polyanthos and
var. rivularis and Chiloscyphus pallescens var. pallescens and var. fragilis). Konstantinova, Bakalin, et al. (2009) also used this concept as well as the World Checklist
of Hornworts and Liverworts (Söderström, Hagborg, et al. 2016). However, several recent publications do not recognise any varieties of the two species (e.g.
Köckinger 2017). The distribution of the taxa recognised here is therefore not recorded consistently over the continent.
82
Heteroscyphus fissitipus is a neophyte in Europe, introduced from Australia/New Zealand where it occurs in three varieties, var. fissistipus, var. multispinus
(E.A.Hodgs. & Allison) J.J.Engel and var. repandus J.J.Engel. Which of these varieties occur in Europe is unknown.
83
Leptoscyphus porphyrius subsp. porphyrius occurs in South America.
84
The Lophocolea bidentata complex has been treated in various ways in the past, and there is also some nomenclatural confusion. Váňa and Engel (2013)
clarify the concepts, not only for Europe but world-wide. As we follow the World Check List (Söderström, Hagborg, et al. 2016), we here treat them in the
genus Lophocolea as Lophocolea bidentata (autoicious) and Lophocolea coadunata (dioicious). Many earlier reports are confusing and the occurrences in
some countries are still questioned.
85
Lophocolea brookwoodiana was described from Britain by Paton and Sheahan (2006).
86
Lophocolea fragrans subsp. cocosana G.Dauphin, Gradst. & M.I.Morales occurs on Cocos Island.
87
Lophocolea heterophylla subsp. cladogyna R.M.Schust. occurs in North America.
88
Lophocolea semiteres var. retusa (J.J.Engel) L.Söderstr. occurs in Australia.
89
Plagiochila bifaria var. rosea (R.M.Schust.) Heinrichs occurs in South America.
90
The primarily neotropical Plagiochila papillifolia was reported as new from the Azores by Heinrichs, Rycroft, et al. (2002).
91
Plagiochila heterophylla var. beauverdii (Steph.) Heinrichs occurs in South America.
92
Plagiochila arctica var. intermedia R.M.Schust. occurs in North America. The synonymy of Plagiochila asplenioides var. lobata (Damsholt 2002) is problematic,
as he states that Plagiochila arctica does not occur in Norway and the type of the synonymised variety is from the mountains in central Norway.
JOURNAL OF BRYOLOGY
10
P. asplenioides (L.) Dumort. [Plagiochila
major (Nees) S.W.Arnell]
11 P. britannica Paton
12 P. porelloides (Torr. ex Nees) Lindenb. [Plagiochila asplenioides subsp. porelloides
(Torr. ex Nees) Kaal.]93
a var. norvegica (H.H.Blom & Holten) Schumacker & Váňa [Plagiochila norvegica H.H.Blom & Holten]
b var. porelloides [Plagiochila asplenioides
var. devexa Carrington, Plagiochila
asplenioides var. minor Lindenb.,
Plagiochila
asplenioides
var.
humilis (Nees) Lindenb.]
c var. subarctica (Jørg.) Lammes [Plagiochila
asplenioides var. subarctica Jørg.]
section Poeltiae Inoue
13 P. carringtonii (Balf. ex Carrington) Grolle
a subsp. carringtonii94
section Rutilantes Carl
14 P. exigua (Taylor) Taylor
15 P. maderensis Gottsche ex Steph.95
section Vagae Lindenb.
16 P. virginica A.Evans [Plagiochila dubia auct.
eur. non Lindenb. & Gottsche (=Plagiochila patula (Sw.) Lindenb.)]
a var. virginica96
Trichocoleaceae Nakai
75 Trichocolea Dumort.
1 T. tomentella (Ehrh.) Dumort.
Myliaceae Schljakov
76 Mylia Gray
subgenus Anomalae (R.M.Schust. ex Potemkin)
L. Söderstr.
1 M. anomala (Hook.) Gray
subgenus Mylia
2 M. taylorii (Hook.) Gray
93
21
Porellales Schljakov
Frullaniaceae Lorch
77
Frullania Raddi
subgenus Frullania
1 F. azorica Sim-Sim, Sérgio, Mues & Kraut
2 F. dilatata (L.) Dumort.
a subsp. dilatata97
3 F. ericoides (Nees) Mont.
a var. ericoides98
4 F. fragilifolia (Taylor) Gottsche, Lindenb. &
Nees
5 F. jackii Gottsche [Frullania davurica subsp.
jackii (Gottsche) S.Hatt.]
6 F. oakesiana Austin
a subsp. oakesiana99
7 F. parvistipula Steph.
8 F. riparia Hampe [Frullania cesatiana De
Not.]100
9 F. stylifera (R.M.Schust.) R.M.Schust.101
subgenus Thyopsiella Spruce
10 F. acicularis Hentschel & von Konrat
[Frullania tamarisci var. azorica J.P.Frahm]102
11 F. calcarifera Steph.103
12 F. microphylla
(Gottsche)
Pearson
[Frullania microphylla var. deciduifolia
Grolle]104
13 F. polysticta Lindenb.
14 F. sergiae Sim-Sim, Fontinha, Mues & Lion
15 F. subarctica Vilnet, Borovich. & Bakalin105
16 F. tamarisci (L.) Dumort. [Frullania tamarisci
var. atrovirens Carrington, Frullania
tamarisci var. cornubica Carrington,
Frullania tamarisci var. ericetorum
Jørg., Frullania tamarisci var. robusta
Lindb., Frullania tamarisci var.
sardoa (De Not.) De Not.]
17 F. teneriffae (F.Weber) Nees
Plagiochila porelloides is a variable species and many varieties have been described. Their value is questionable but the three included here have frequently
been recognised in recent studies.
Plagiochila carringtonii subsp. lobuchensis Grolle occurs in Himalaya.
95
Plagiochila maderensis was placed in synonymy with Plagiochila spinulosa by Grolle (1967) but reinstated by Rycroft et al. (2004).
96
Plagiochila virginica var. caroliniana R.M.Schust. and var. euryphylla R.M.Schust. occur in North America.
97
Frullania dilatata subsp. asiatica S.Hatt. occurs in E Asia.
98
Frullania ericoides var. laxa (Gottsche, Lindenb. & Nees) Schiffn. occurs in SE Asia and Mexico, var. minor Kamim. in Japan and var. verrucosa (Kamim.)
Hentschel & von Konrat in Japan.
99
Frullania oakesiana subsp. takayuensis (Steph.) R.M.Schust. occurs in E Asia.
100
In spite of being under discussion for over 30 years, the problem of the Frullania riparia (American) / cesatiana (European) / muscicola (Asiatic) complex has
not yet been solved. As concluded in Bisang et al. (1989), if they are the same, the name Frullania riparia has priority. However, if the American populations
are different from the European and Asiatic populations (as treated in Bisang et al. 1989), Frullania cesatiana would be the correct name (note that their
‘combination’ Frullania cesatiana var. muscicola (Steph.) Bisang et al. is a nomen nudum). Grolle (in Grolle and Long 2000) argued that the American Frullania riparia and European Frullania cesatiana are conspecific, but separate from the Asiatic Frullania muscicola Steph. For now we are following this
concept pending further research, preferably including molecular data.
101
Frullania stylifera, until recently considered endemic to eastern North America, has recently been recognised from European and Asian Russia (Konstantinova et al. 2020).
102
Frullania acicularis was described from the Azores by Frahm (2006) as Frullania tamarisci var. azorica but shown to be separate from Frullania tamarisci by
Heinrichs, Hentschel, et al. (2010) and Vilnet, Borovichev, et al. (2014) and elevated to species level by Hentschel et al. (2015)
103
Frullania calcarifera was placed in synonymy of Frullania tamarisci by Hattori (1972) but shown to be a species separate from it by Heinrichs, Hentschel,
et al. (2010) and Vilnet, Borovichev, et al. (2014).
104
Frullania microphylla var. deciduifolia is not recognized by Sim-Sim (1999) as “all specimens from Portugal or Madeira have deciduous leaves”’, a treatment
followed here as well as in the world checklist (Söderström, Hagborg, et al. 2016).
105
Frullania subarctica was described by Vilnet, Borovichev, et al. (2014) from the Russian Far East but shown also to occur in northern Europe.
94
22
N. G. HODGETTS ET AL.
7 C. azorica V.Allorge & Jovet-Ast [Aphanolejeunea azorica (V.Allorge & Ast) Bernecker
& Pócs]
incertae sedis
18 F. bolanderi Austin
19 F. cleistostoma Schiffn. & W.Wollny [Frullania
inflata auct. eur. non Gottsche]106
82
Jubulaceae H.Klinggr.
78 Jubula Dumort.
1 J. hutchinsiae (Hook.) Dumort.107
a subsp. caucasica Konstant. & Vilnet108
b subsp. hutchinsiae
Colura (Dumort.) Dumort.
subgenus Colura
section Colura
1 C. calyptrifolia (Hook.) Dumort.
83
Lejeuneaceae Cavers
Lejeuneoideae C.Massal.
trib. Brachiolejeuneeae van Slageren & Berendsen
subtrib. Brachiolejeuneinae Gradst.
79 Acanthocoleus R.M.Schust.
1 A. aberrans (Lindenb. & Gottsche) Kruijt
a var. laevis Gradst.109
Myriocoleopsis Schiffn.
1 M. minutissima (Sm.) R.L.Zhu, Y.Yu & Pócs [Cololejeunea minutissima (Sm.) Schiffn.]
a subsp. minutissima111
subtrib. Drepanolejeuneinae Gradst.
84 Drepanolejeunea (Spruce) Steph.
subgenus Drepanolejeunea
1 D. hamatifolia (Hook.) Schiffn.
trib. Lejeuneeae Dumort.
subtrib. Cheilolejeuneinae Gradst.
80 Cheilolejeunea (Spruce) Steph.
subgenus Euosmolejeunea (Spruce) Kachroo
1 C. cedercreutzii (H.Buch & Perss.) Grolle
subtrib. Lejeuneinae Gradst.
85 Harpalejeunea (Spruce) Schiffn.
subgenus Harpalejeunea
1 H. molleri (Steph.) Grolle
a subsp. molleri112
subtrib. Cololejeuneinae Gradst.
81 Cololejeunea (Spruce) Steph. [Aphanolejeunea
A.Evans]
subgenus Aphanolejeunea (A.Evans) Pócs
1 C. madeirensis Schiffn. [Aphanolejeunea
madeirensis (Schiffn.) Grolle]
2 C. microscopica (Taylor) Schiffn. [Aphanolejeunea microscopica (Taylor) A.Evans]
a var. microscopica110
3 C. sintenisii (Steph.) Pócs [Aphanolejeunea sintenisii Steph.]
subgenus Cololejeunea
4 C. calcarea (Lib.) Steph.
5 C. rossettiana (C.Massal.) Schiffn.
6 C. schaeferi Grolle
subgenus Diaphanae R.M.Schust.
86
106
Lejeunea Lib.
subgenus Lejeunea
1 L. cavifolia (Ehrh.) Lindb.
2 L. eckloniana Lindenb. [Lejeunea holtii
Spruce]113
3 L. flava (Sw.) Nees
a subsp. moorei (Lindb.) R.M.Schust.114
4 L. hibernica Bischl., H.A.Mill. & Bonner ex
Grolle
5 L. lamacerina (Steph.) Schiffn.
a subsp. lamacerina115
6 L. mandonii (Steph.) Müll.Frib.
7 L. patens Lindb.
subgenus Nanolejeunea R.M.Schust.
8 L. canariensis (Steph.) Steph. [Lejeunea laetevirens auct.]116
Frullania inflata Gottsche as treated in Söderström, Hagborg, et al. (2016) is a species complex. Frullania inflata s.str. is confined to North America and
reports from Europe belongs to the segregate Frullania cleistostoma (Mamontov, Potemkin, et al. 2018).
107
Jubula hutchinsiae is a complex species with many subspecies. In addition to the two subspecies occurring in Europe, five more are recognised in Söderström, Hagborg, et al. (2016), subsp. australiae Pócs & A.Cairns (Australia), subsp. bogotensis (Steph.) Verd. (S and C America), subsp. japonica (Steph.) Horik.
& Ando (E Asia), subsp. javanica (Steph.) Verd. (E and SE Asia) and subsp. pennsylvanica (Steph.) Verd. (North America).
108
Jubula hutchinsiae subsp. caucasica was described by Konstantinova and Vilnet (2011). Prior to the description, the subspecies occurring in the Caucasus
was regarded as subsp. javanica (Steph.) Verd.
109
Acanthocoleus aberrans var. aberrans seems to be confined to South America.
110
Cololejeunea microscopica var. africana (Pócs) Pócs & Bernecker and var. exigua (A.Evans) Pócs occur in Africa and South America.
111
Myriocoleopsis minutissima subsp. myriocarpa (Nees & Mont.) R.L.Zhu, Y.Yu & Pócs is widespread in the tropics.
112
Harpalejeunea molleri subsp. integra (R.M.Schust.) Damsh. occurs in North America.
113
Lejeunea holtii was placed in synonymy with Lejeunea eckloniana by Dirkse et al. (1993). However, the two taxa differ slightly in morphology and a molecular study is needed to verify the synonymy.
114
Lejeunea flava subsp. flava is widespread in the tropics, subsp. orientalis R.M.Schust. occurs in E and SE Asia, subsp. tabularis (Spreng.) S.W.Arnell in Africa
and var. pellucida Lindenb. & Gottsche in South America.
115
Lejeunea lamacerina subsp. gemminata R.M.Schust. occurs in North America.
116
Lejeunea canariensis from Madeira was nested within Lejeunea laetevirens in the study by Heinrichs, Dong, et al. (2013). However, Lejeunea laetevirens
seems to be a complex species with several other taxa also nested within it. We keep Lejeunea canariensis as a separate species for now, recognizing
that it seems to be the only element of the Lejeunea laetevirens complex occurring in Macaronesia.
JOURNAL OF BRYOLOGY
87
6 R. visianica C.Massal.118
7 R. wichurae Steph.
subgenus Volutoradula Devos, M.A.M.Renner,
Gradst., A.J.Shaw & Vanderp.
8 R. holtii Spruce
9 R. nudicaulis Steph.
a var. delicatula P.Allorge & V.Allorge
b var. nudicaulis
10 R. voluta Taylor
Microlejeunea (Spruce) Steph.
1 M. ulicina (Taylor) Steph. [Lejeunea ulicina
(Taylor) Gottsche, Lindenb. & Nees]
Ptychanthoideae Mizut.
88 Marchesinia Gray
subgenus Marchesinia
1 M. mackaii (Hook.) Gray
Porellaceae Cavers
89 Porella L.
1 P. arboris-vitae (With.) Grolle
a subsp. arboris-vitae [Porella arboris-vitae
var. killarniensis (Pearson) M.F.V.Corley, Porella arboris-vitae var. obscura
(Nees) M.F.V.Corley]117
2 P. baueri (Schiffn.) C.E.O.Jensen
3 P. canariensis (F.Weber) Underw.
4 P. cordaeana (Huebener) Moore [Porella cordaeana var. faeroensis (C.E.O.Jensen)
E.W.Jones, Porella cordaeana var. simplicior (J.E.Zetterst.) Arnell]
5 P. inaequalis (Gottsche) Perss.
6 P. obtusata (Taylor) Trevis.
7 P. pinnata L.
8 P. platyphylla (L.) Pfeiff. [Porella platyphylla
var. subsquarrosa (Schiffn.) Arnell]
Radulaceae Müll.Frib.
90 Radula Dumort.
subgenus Radula
1 R. aquilegia (Hook.f. & Taylor) Gottsche,
Lindenb. & Nees
2 R. carringtonii J.B.Jack
3 R. complanata (L.) Dumort. [Radula complanata var. alpestris (Lindb. ex Berggr.)
Lindb.]
4 R. jonesii Bouman, Dirkse & K.Yamada
5 R. lindenbergiana Gottsche ex C.Hartm.
[Radula complanata subsp. lindenbergiana (Gottsche ex C.Hartm.)
R.M.Schust.,
Radula
lindbergiana
Gottsche ex J.B.Jack nom. inval.]
117
23
Ptilidiales Schljakov
Ptilidiaceae H.Klinggr.
91 Ptilidium Nees
1 P. ciliare (L.) Hampe
2 P. pulcherrimum (Weber) Vain. [Ptilidium pulcherrimum var. subpinnatum (Jørg.) Damsh.]
Metzgeriales Chalaud
Aneuraceae H.Klinggr.
92 Aneura Dumort [Cryptothallus Malmb.]119
1 A. latissima Spruce [Aneura pseudopinguis
(Herzog) Pócs]120
2 A. maxima (Schiffn.) Steph.121
3 A. mirabilis (Malmb.) Wickett & Goffinet [Cryptothallus mirabilis Malmb.]
4 A. pinguis (L.) Dumort. [Aneura pinguis var. angustior (Hook.) Dumort., Aneura pinguis var.
denticulata (Nees) Godelin., Aneura
pinguis var. fuscovirens (Lindb.) Damsh.]
93
Riccardia Gray122
1 R. chamedryfolia (With.) Grolle [Riccardia chamedryfolia
var.
major
(Nees)
R.M.Schust. nom. illeg., Riccardia chamedryfolia var. submersa (C.E.O.Jensen
ex Müll.Frib.) Damsh. nom. inval.]
2 R. incurvata Lindb.
3 R. latifrons (Lindb.) Lindb.
a subsp. arctica R.M.Schust. & Damsh.
b subsp. latifrons123
4 R. multifida (L.) Gray
a subsp. multifida124
5 R. palmata (Hedw.) Carruth.
Porella arboris-vitae subsp. nitidula (C.Massal.) S.Hatt. occurs in E Asia.
Radula visianica was until recently only known from two collections in N. Italy, the type in 1878 and a later collection in the southern Alps in the 1930s, but
was recently found in several localities in Austria (Köckinger 2016). It has been shown to be well separated molecularly from other species in subgenus
Radula (Váňa et al. 2017).
119
The genus Aneura includes several genetically distinct lineages (cf. Bączkiewicz et al. 2017) and the European ‘Aneura maxima’ is genetically different from
the population in the type locality on Java (D.G.Long, pers. comm.). It is also unlikely that Aneura latissima (syn. Aneura pseudopinguis) from South America
represents any of the European lineages. However, until the taxonomy is further clarified, the ‘traditional’ treatment is followed here, as it was in Söderström, Hagborg, et al. (2016).
120
Aneura latissima was reported (as Aneura pseudopinguis) new to Europe by Sérgio and Garcia (2009). The type of both names is from South America and it
is unlikely that the European plants represent this taxon. It is more probable that they belong to some of the unpublished European lineages (Long et al. in
prep.).
121
Aneura maxima is a SE Asian taxon that is not conspecific with the taxon in Europe so named. The latter is at present lacking a valid name.
122
Riccardia is classified into several subgenera and sections in Söderström, Hagborg, et al. (2016). The European species all belong to subgenus Riccardia, and
mostly also to section Riccardia, except for R. incurvata, the position of which is unclear.
123
Riccardia latifrons var. miyakeana (Schiffn.) Furuki occurs in Japan.
124
Riccardia multifida subsp. decrescens (Steph.) Furuki occurs in E Asia and subsp. synoica R.M.Schust. in SE USA.
118
24
N. G. HODGETTS ET AL.
Metzgeriaceae H.Klinggr.
94 Metzgeria Raddi [Apometzgeria Kuwah.]
1 M. conjugata Lindb. [Metzgeria conjugata var.
alipila Kaal., Metzgeria conjugata var.
macvicarii Kaal.]
2 M. consanguinea Schiffn. [Metzgeria temperata auct. eur. non Kuwah.]125
3 M. furcata (L.) Corda [Metzgeria furcata var.
expansa Douin, Metzgeria furcata var.
flexipilis Kaal., Metzgeria furcata var.
ulvula (Nees) Pavletic]126
4 M. leptoneura Spruce [Metzgeria hamata auct.
non Lindb.]127
a var. leptoneura128
5 M. pubescens (Schrank) Raddi [Apometzgeria
pubescens (Schrank) Kuwah.]
6 M. simplex Lorb. ex Müll.Frib. [Metzgeria conjugata subsp. simplex (Lorb. ex Müll.Frib.)
R.M.Schust.]
7 M. violacea (Ach.) Dumort. [Metzgeria fruticulosa auct. non (O.F.Müll.) A.Evans]129
Pleuroziales Schljakov
Pleuroziaceae Müll.Frib.
95 Pleurozia Dumort.
subgenus Constantifolia B.M.Thiers
1 P. purpurea Lindb.
Fossombroniales Schljakov
Calyculariaceae He-Nygrén, Juslén, Ahonen, Glenny &
Piippo
96 Calycularia Mitt.
1 C. laxa Lindb. & Arnell130
Fossombroniaceae Hazsl.
97 Fossombronia Raddi
1 F. angulosa (Dicks.) Raddi
2 F. caespitiformis (Raddi) De Not. ex Rabenh.
125
3
4
5
6
7
8
9
10
11
12
13
a subsp. caespitiformis
b subsp. multispira (Schiffn.) J.R.Bray &
Cargill [Fossombronia husnotii
Corb.]
F. crispa Nees [Fossombronia zeyheri
Steph.]131
F. echinata Macvicar
F. fimbriata Paton
F. fleischeri Osterwald ex Loeske132
F. foveolata Lindb.
F. incurva Lindb.
F. leucoxantha (Lehm.) Lehm. & Lindenb.133
F. maritima (Paton) Paton
F. mittenii Tind. [Fossombronia crozalsii Corb.]
F. pusilla (L.) Nees
F. wondraczekii (Corda) Dumort. ex Lindb.
Petalophyllaceae Stotler & Crand.-Stotl.
98 Petalophyllum Nees & Gottsche
1 P. ralfsii (Wilson) Nees & Gottsche
Pallaviciniales W.Frey & M.Stech
Moerckiaceae K.I.Goebel ex Stotler & Crand.-Stotl.
99 Moerckia Gottsche
1 M. blyttii (Mørch) Brockm.
2 M. flotoviana (Nees) Schiffn.134
3 M. hibernica (Hook.) Gottsche
Pallaviciniaceae Mig.
Pallavicinioideae Mig. ex Grolle
100 Pallavicinia Gray
1 P. lyellii (Hook.) Gray
Pelliales He-Nygrén
Pelliaceae H.Klinggr.
101 Apopellia (Grolle) Nebel & D.Quandt
1 A. endiviifolia (Dicks.) Nebel & D.Quandt
[Pellia endiviifolia (Dicks.) Dumort.]
Metzgeria temperata was synonymised with the primarily pantropical Metzgeria consanguinea by Hill et al. (2008). However, unpublished molecular studies
show that the European populations are closer to Metzgeria violacea than to the Japanese Metzgeria temperata (see Köckinger 2017). Until the position of
the European Metzgeria ‘temperata’ is clarified, we use the name Metzgeria consanguinea, although with some hesitation.
126
North American populations of Metzgeria furcata have recently been recognised as a separate species, Metzgeria setigera R.M.Schust. ex Crand.-Stotl. &
L. Söderstr. (Söderström, Váňa, et al. 2015). Metzgeria furcata var. pacifica Brinkm. occurs in E North America, and probably belongs to Metzgeria setigera.
Metzgeria furcata var. expansa was recognised in Söderström, Hagborg, et al. (2016), although with low confidence. It is known only from the type specimen in France and we do not recognise it here, as there are several other equally doubtful ’varieties‘ that could also be included. The whole Metzgeria
furcata complex needs further study in Europe (and elsewhere).
127
Metzgeria hamata Lindb. is technically a synonym of Metzgeria procera Mitt., but identical to Metzgeria leptoneura in the sense of Lindberg (1877) and all
subsequent authors.
128
Metzgeria leptoneura var. breviseta (Schiffn.) O.Yano occurs in Brazil and var. polychaeta R.M.Schust. in SE USA.
129
Grolle and So (2003) demonstrated that the name Metzgeria fruticulosa technically belongs to Riccardia palmata and that Metzgeria violacea is the name
that should be used.
130
Calycularia laxa was reported new to European Russia from Nenets (Konstantinova and Lavrinenko 2002) and Murmansk (Sofronova et al. 2013) provinces.
131
Fossombronia crispa was first reported from Portugal by Sérgio (1985) but Perold (1997) showed that her concept of the species was wrong. Later, Sérgio
(2003) referred the plants to Fossombronia leucoxantha. However, Sérgio (1985) also reported Fossombronia zeyheri, which Perold (1997) showed was a
synonym of Fossombronia crispa but Grolle and Long (2000) treated it as a synonym of Fossombronia foveolata.
132
Fossombronia fleischeri has long been considered a synonym of Fossombronia incurva, but Stotler et al. (2003) showed that they should be regarded as
distinct species.
133
Fossombronia leucoxantha was reported from Portugal by Sérgio (1985) as Fossombronia crispa but Perold (1997) pointed out that her concept of Fossombronia crispa was wrong. Sérgio (2003) later referred the plants to Fossombronia leucoxantha.
134
Moerckia flotoviana was synonymised with Moerckia hibernica by De Sloover (1959) and subsequently usually treated as such in the European literature.
However, Crandall-Stotler and Stotler (2007) showed that they are distinct taxa. Thus, many reports of Moerckia hibernica belong to Moerckia flotoviana.
JOURNAL OF BRYOLOGY
102
Pellia Raddi
1 P. epiphylla (L.) Corda
a subsp. borealis (Lorb.) Messe
b subsp. epiphylla
2 P. neesiana (Gottsche) Limpr.
MARCHANTIOPSIDA CRONQUIST , TAKHT . & W.ZIMM .
Blasiales Stotler & Crand.-Stotl.
Blasiaceae H.Klinggr.
103 Blasia L.
1 B. pusilla L.
Lunulariales H.Klinggr.
Lunulariaceae H.Klinggr.
104 Lunularia Adans.
1 L. cruciata (L.) Dumort. ex Lindb.
a subsp. cruciata135
Marchantiales Limpr.
Aytoniaceae Cavers
105 Asterella P.Beauv. [Fimbraria Nees]
subgenus Asterella
section Brachyblepharis (Nees) D.G.Long
1 A. africana (Mont.) Underw. ex A.Evans
subgenus Phragmoblepharis Grolle
2 A. lindenbergiana (Corda ex Nees) Lindb. ex
Arnell
subgenus Saccatae (Grolle) D.G.Long
3 A. saccata (Wahlenb.) A.Evans
106
135
Mannia Corda
subgenus Mannia
1 M. androgyna (L.) A.Evans
2 M. californica (Gottsche) L.C.Wheeler136
3 M. controversa (Meyl.) D.B.Schill137
a subsp. controversa138
4 M. fragrans (Balb.) Frye & L.Clark
a subsp. fragrans [Mannia fragrans var. brevipes (Kaal.) Damsh.]139
5 M. sibirica (Müll.Frib.) Frye & L.Clark
subgenus Neesiella
6 M. gracilis (F.Weber) D.B.Schill & D.G.Long
[Asterella gracilis (F.Weber) Underw.]
25
7 M. pilosa (Hornem.) Frye & L.Clark
8 M. triandra (Scop.) Grolle
107
Plagiochasma Lehm.
subgenus Micropylum
1 P. rupestre (J.R.Forst. & G.Forst.) Steph.
a var. rupestre140
subgenus Plagiochasma
2 P. appendiculatum Lehm. & Lindenb.141
108
Reboulia Raddi
1 R. hemisphaerica (L.) Raddi
a subsp. australis R.M.Schust.
b subsp. dioica R.M.Schust.
c subsp. hemisphaerica142
Cleveaceae Cavers
109 Clevea Lindb. [Athalamia auct. eur.]
1 C. hyalina (Sommerf.) Lindb. [Athalamia
hyalina (Sommerf.) S.Hatt., Clevea
hyalina var. suecica (Lindb.) Lindb.,
Athalamia hyalina var. suecica (Lindb.)
S.Hatt., Clevea hyalina var. rufescens
(S.W.Arnell) Konstant. nom. inval.]
2 C. spathysii (Lindenb.) Müll.Frib. [Athalamia
spathysii (Lindenb.) S.Hatt.]
110
Peltolepis Lindb.
1 P. quadrata (Saut.) Müll.Frib.
111
Sauteria Nees
Section Sauteria
1 S. alpina (Nees) Nees
Conocephalaceae Müll.Frib. ex Grolle
112 Conocephalum Hill
1 C. conicum (L.) Dumort.
2 C. salebrosum Szweyk., Buczk. & Odrzyk.143
Corsiniaceae Engl.
Corsinioideae Schiffn.
113 Corsinia Raddi
1 C. coriandrina (Spreng.) Lindb.
Lunularia cruciata subsp. thaxteri (A.Evans & Herzog) R.M.Schust. occurs in South America and New Zealand.
Mannia californica was reported new to Europe by Hugonnot and Schill (2006).
137
Mannia controversa (type from Switzerland) was usually regarded as a synonym of Mannia fragrans until Schill et al. (2008) showed it to be an independent
species.
138
Mannia controversa subsp. asiatica D.B.Schill & D.G.Long occurs in Asia.
139
Mannia fragrans subsp. orientalis R.M.Schust. occurs in Japan.
140
Plagiochasma rupestre var. volkii Bischl. occurs in South America and southern Africa.
141
Plagiochasma appendiculatum was reported new to Europe from the Baleares by Cros et al. (2005).
142
Reboulia hemispherica subsp. paradoxa R.M.Schust. is described from Portugal (Schuster 1992b) but invalidly as it lacks a Latin description. If recognised it
needs to be validated, but there are many uncertainties about the segregates of Reboulia hemispherica. Reboulia hemispherica subsp. acrogyna
(R.M.Schust.) R.M.Schust. occurs in North America, subsp. orientalis R.M.Schust. in Asia, var. fissisquama Herzog in China and var. turkestanica C.E.O.Jensen
ex Herzog in China.
143
Conocephalum salebrosum was described by Szweykowski et al. (2005) from Poland but shown to have a wide distribution both in Europe and elsewhere in
the Northern Hemisphere. On the other hand, they showed that Conocephalum conicum was a near-endemic to Europe.
136
26
N. G. HODGETTS ET AL.
Cyathodiaceae Stotler & Crand.-Stotl.
114 Cyathodium Kunze
1 C. foetidissimum Schiffn.144
Dumortieraceae D.G.Long
115
Dumortiera Nees
1 D. hirsuta (Sw.) Nees
a subsp. hirsuta145
Exormothecaceae Müll.Frib. ex Grolle
116 Exormotheca Mitt.
subgenus Corbierella (Douin & Trab.) Schiffn.
1 E. welwitschii Steph.
subgenus Exormotheca
2 E. pustulosa Mitt.
Marchantiaceae Lindl.
Marchantioideae Schiffn.
117
Marchantia L.
subgenus Chlamidium (Corda) Bischl.
section Paleacea Bischl.
1 M. paleacea Bertol.
a subsp. paleacea146
subgenus Marchantia
2 M. polymorpha L.147
a subsp. montivagans Bischl. & Boissel.Dub. [Marchantia alpestris (Nees)
Burgeff]
b subsp. polymorpha [Marchantia aquatica
(Nees) Burgeff]
c subsp. ruderalis Bischl. & Boissel.-Dub.
[Marchantia latifolia Gray]
subgenus Preissia (Corda) D.G.Long, Crand.-Stotl.,
L.L.Forrest & Villarreal [Bucegia
Radian, Preissia Corda]
3 M. quadrata Scop. [Preissia quadrata (Scop.)
Nees]
a subsp. hyperborea (R.M.Schust.) Borovich. [Preissia quadrata subsp.
hyperborea R.M.Schust.]
b subsp. quadrata
4 M. romanica (Radian) D.G.Long, Crand.-Stotl.,
L.L.Forrest & J.C.Villarreal [Bucegia
romanica Radian]
Oxymitraceae Müll.Frib. ex Grolle
144
118
Oxymitra Bisch. ex Lindenb.
1 O. incrassata (Brot.) Sérgio & Sim-Sim
Ricciaceae Rchb.
119 Riccia L.
subgenus Riccia
section Riccia148
1 R. atlantica Sérgio & Perold
2 R. atromarginata Levier
a var. atromarginata149
3 R. beyrichiana Hampe [Riccia marginata Lindb.]
4 R. bicarinata Lindb.
5 R. bifurca Hoffm. [Riccia bifurca var. subinermis Heeg]
6 R. breidleri Jur. ex Steph.
7 R. ciliata Hoffm. [Riccia ciliata var. epilosa
Warnst., Riccia ciliata var. intumescens
Bisch., Riccia ciliata var. violacea Kny,
Riccia dalslandica S.W.Arnell, Riccia
intumescens (Bisch.) Underw., Riccia
canescens Steph., Riccia trichocarpa
M.Howe, Riccia crinita auct. eur.]150
8 R. ciliifera Link [Riccia melitensis C.Massal.]
9 R. crozalsii Levier
10 R. crustata Trab.
11 R. glauca L. [Riccia glauca var. major (Roth)
Lindenb.]
a var. ciliaris Warnst. [Riccia glauca var.
subinermis (Lindb.) Warnst.]
b var. glauca
12 R. gothica Damsh. & Hallingb.
13 R. gougetiana Durieu & Mont.
a var. armatissima Levier ex Müll.Frib.
b var. gougetiana
14 R. lamellosa Raddi
15 R. ligula Steph.
16 R. macrocarpa Levier
17 R. michelii Raddi
18 R. nigrella DC.
19 R. papillosa Moris
20 R. sommieri Levier
21 R. sorocarpa Bisch.
a subsp. arctica R.M.Schust. ex Köckinger
& L. Söderstr. [Riccia sorocarpa
subsp. arctica R.M.Schust. nom.
inval., Riccia lindenbergiana Saut.]
Cyathodium foetidissimum was recorded new to Europe by Ligrone and Duckett (2005).
Dumortiera hirsuta subsp. nepalensis (Taylor) R.M.Schust. is widespread in the tropics and subsp. tatunoi Horik. occurs in Japan.
146
Marchantia paleacea subsp. diptera (Nees & Mont.) Inoue occurs in E Asia.
147
Recent molecular studies (Cronberg et al., unpubl.) indicate that the subspecies of Marchantia polymorpha should be better treated at species level.
However, we keep them as subspecies until nomenclatural uncertainties raised by elevating them to species are resolved.
148
As noted by Hugonnot (2010a), the nomenclature and taxonomy of many species in sect. Riccia is highly problematic. Difficult taxa include Riccia bicarinata, R. bifurca, R. ciliata, R. crozalsii, R. michelii, R. subbifurca, R. warnstorfii and the little-known R. ligula. The morphology of these taxa is plastic, and it is
unclear to what extent the size of the thallus and the development and structure of cilia are determined by genetic or environmental factors. Jovet-Ast
(1986) published an important monograph of the Mediterranean species, but her concepts have been questioned in recent years (e.g. Hugonnot 2010b,
2015). Integrated molecular and morphological studies are urgently needed.
149
Riccia atromarginata var. jovet-astiae Rauh & Buchloh occurs on the Arabian Peninsula and adjacent parts of Africa.
150
Riccia trichocarpa was synonymised with the Australian Riccia crinita (the latter having priority) by Jovet-Ast (2000), a synonymisation that has been largely
overlooked or rejected by European bryologists. Hugonnot (2010b) argued that Jovet-Ast’s (1986) concept of Riccia trichocarpa is actually Riccia ciliata, and
her Riccia ciliata belongs to other species. Thus Hugonnot synonymised both taxa under the oldest name, Riccia ciliata.
145
JOURNAL OF BRYOLOGY
b
subsp. erythrophora R.M.Schust. ex
Konstant. & L.Söderstr. [Riccia
sorocarpa subsp. erythrophora
R.M.Schust. nom. inval.]
c subsp. sorocarpa151 [Riccia sorocarpa
var. heegii Schiffn.]
22 R. subbifurca Warnst. ex Croz. [?Riccia oelandica C.E.O.Jensen]152
23 R. trabutiana Steph.
24 R. warnstorfii Limpr. ex Warnst. [Riccia
warnstorfii var. commutata (J.B.Jack
ex Levier) Damsh., Riccia warnstorfii
var. subinermis Warnst., Riccia warnstorfii var. ciliaris Warnst.]
section Pilifer O.H.Volk
25 R. boumanii Dirkse, Losada-Lima & M.Stech153
subgenus Ricciella (A.Braun) Boulay
section Ricciella (A.Braun) Bisch.
26 R. canaliculata Hoffm.
27 R. duplex Lorb. ex Müll.Frib.
a var. duplex154
28 R. fluitans L.
29 R. frostii Austin
a var. frostii155
30 R. huebeneriana Lindenb.
a subsp. huebeneriana [Riccia huebeneriana var. pseudo-frostii Schiffn.]156
31 R. perennis Steph.
32 R. rhenana Lorb. ex Müll.Frib.
a var. rhenana
b var. violacea M.F.Boiko
section Spongodes Nees
33 R. cavernosa Hoffm. [Riccia teneriffae
S.W.Arnell, Riccia cavernosa var. angustior (Nees) Damsh.]
34 R. crystallina L.
120
Ricciocarpos Corda
1 R. natans (L.) Corda
Targioniaceae Dumort.
121
151
27
subgenus Targionia
1 T. hypophylla L.
a subsp. hypophylla157
2 T. lorbeeriana Müll.Frib.
Sphaerocarpales Cavers
Riellaceae Engl.
122 Riella Mont.
subgenus Riella
1 R. battandieri Trab.
2 R. bialata Trab.158
3 R. gallica Balansa ex Trab.
4 R. helicophylla (Bory & Mont.) Mont.
a var. helicophylla
b var. macrocarpa P.Allorge
5 R. notarisii (Mont.) Mont.159
6 R. reuteri Mont.
subgenus Trabutiella Porsild
7 R. affinis M.Howe & Underw.
8 R. cossoniana Trab.
9 R. echinata (Müll.Frib.) Segarra, Puche &
Sabovlj.160
10 R. mediterranea Segarra, Puche, Sabovlj.,
M.Infante & Heras161
Sphaerocarpaceae Heeg
123 Sphaerocarpos Boehm.
subgenus Austrosphaerocarpos R.M.Schust.
1 S. stipitatus Bisch. ex Lindenb.
subgenus Sphaerocarpos
2 S. europaeus Lorb. [Sphaerocarpos texanus
auct. eur. non Austin]162
3 S. michelii Bellardi
Mosses
BRYOPHYTA
SPHAGNOPSIDA SCHIMP .
Sphagnales Limpr.
Sphagnaceae Dumort.
1 Sphagnum L.163
Targionia L.
Riccia sorocarpa var heegii was accepted with low confidence in the world checklist of liverworts (Söderström, Hagborg, et al. 2016). In Europe it has been
reported from Portugal, Spain, France, Corsica, Sardinia, Austria, Serbia, North Macedonia and Romania as well as from the Canary Islands and Madeira.
152
The name Riccia subbifurca has been used for different plants in Europe, and the application of the name is problematic, as is its relationship with other
taxa such as R. oelandica C.E.O. Jensen. The species was described by Crozals (1903), based primarily on material from a serpentine site in France. It is
discussed by Hugonnot (2018), who suggests that it could be a taxon of localised occurrence on serpentine.
153
Riccia boumanii is newly described from the Canary Islands by Dirkse et al. (2016).
154
Riccia duplex var. megaspora Na-Thalang occurs in Australia.
155
Riccia frostii var. crystallinoides Schiffn. occurs in W Asia.
156
Riccia huebeneriana subsp. sullivantii (Austin) R.M.Schust. occurs in North America.
157
Targionia hypophylla subsp. linealis W.Frey & Kürschner occurs on the Arabian peninsula.
158
Riella bialata was reported new to Europe from Spain by Puche and Segarra-Moragues (2013).
159
Riella notarisii is a species complex with poorly defined taxonomic boundaries including the European Riella battandieri, Riella gallica and Riella reuteri as
well as Riella cyrenaica Maire and Riella sersuensis Trab. from North Africa.
160
Riella echinata is a new species described from North Africa by Segarra-Moragues et al. (2014) but also occurring in Spain, The Baleares and Canary Islands.
161
Riella mediterranea is a new species described from Spain by Segarra-Moragues et al. (2014) but also occurring in the Baleares, Malta and Cyprus as well as
in Morocco.
162
European accessions of ‘Sphaerocarpos texanus’ were shown to be clearly distinct from the American Sphaerocarpos texanus (Bell et al. 2013) and the latter
is thus excluded from Europe. Sphaerocarpos europaeus is the oldest name based on a European specimen.
163
The subgeneric and sectional classification of Sphagnum follows Laine et al. (2018).
28
N. G. HODGETTS ET AL.
subgenus Rigida (Lindb.) A.Eddy
1 S. compactum Lam. & DC.
2 S. strictum Sull.
subgenus Sphagnum
3 S. affine Renauld & Cardot [Sphagnum affine var.
flagellare (Schlieph. ex Röll) L.Söderstr. &
Hedenäs, Sphagnum imbricatum subsp.
affine (Renauld & Cardot) Flatberg]
4 S. austinii Sull. [Sphagnum imbricatum subsp.
austinii (Sull.) Flatberg]
5 S. centrale C.E.O.Jensen [Sphagnum palustre
var. centrale (C.E.O.Jensen) A.Eddy]
6 S. divinum Flatberg & Hassel [Sphagnum
magellanicum auct. eur. p.p., non Brid.]
7 S. medium Limpr. [Sphagnum magellanicum
auct. eur. p.p., non Brid.]164
8 S. palustre L.
9 S. papillosum Lindb.
subgenus Acutifolia (Russow) A.J.Shaw
section Squarrosa (Russow) Schimp.
10 S. mirum Flatberg & Thingsgaard165
11 S. squarrosum Crome
12 S. teres (Schimp.) Ångstr.
13 S. tundrae Flatberg
section Polyclada (C.E.O.Jensen) Horrell
14 S. wulfianum Girg.
section Insulosa Isov.
15 S. aongstroemii C.Hartm.
section Acutifolia Wilson
16 S. angermanicum Melin
17 S. arcticum Flatberg & Frisvoll
18 S. beothuk R.E.Andrus166
19 S. capillifolium (Ehrh.) Hedw. [Sphagnum capillifolium subsp. capillifolium]
20 S. concinnum (Berggr.) Flatberg [Sphagnum
fimbriatum subsp. concinnum (Berggr.)
Flatberg & Frisvoll]167
21 S. fimbriatum Wilson
22 S. fuscum (Schimp.) H.Klinggr.
23 S. girgensohnii Russow
24 S. molle Sull.
25 S. nitidulum Warnst.168
26 S. olafii Flatberg
164
27
28
S. quinquefarium (Braithw.) Warnst.
S. rubellum Wilson [Sphagnum capillifolium
subsp. rubellum (Wilson) M.O.Hill]
29 S. rubiginosum Flatberg
30 S. russowii Warnst.
31 S. skyense Flatberg
32 S. subfulvum Sjörs
a subsp. purpureum Flatberg
b subsp. subfulvum
33 S. subnitens Russow & Warnst.
a subsp. ferrugineum Flatberg [Sphagnum
subnitens var. ferrugineum (Flatberg) M.O.Hill]
b subsp. subnitens
34 S. tescorum Flatberg169
35 S. venustum Flatberg170
36 S. warnstorfii Russow
subgenus Subsecunda (Lindb.) A.J.Shaw
37 S. auriculatum Schimp. [Sphagnum denticulatum Brid.]
38 S. contortum Schultz
39 S. inundatum Russow
40 S. platyphyllum (Lindb. ex Braithw.) Warnst.
41 S. pylaesii Brid.
42 S. subsecundum Nees
subgenus Cuspidata Lindb.
43 S. angustifolium (C.E.O.Jensen ex Russow)
C.E.O.Jensen
44 S. annulatum H.Lindb. ex Warnst.
45 S. balticum (Russow) C.E.O.Jensen
46 S. cuspidatum Ehrh. ex Hoffm.
a var. cuspidatum
b var. viride (Flatberg) Lönnell & Hassel
[Sphagnum viride Flatberg]171
47 S. fallax (H.Klinggr.) H.Klinggr.172
a var. brevifolium (Lindb. ex Braithw.)
Lönnell & Hassel [Sphagnum
brevifolium (Lindb. ex Braithw.)
Röll]
b var. fallax
c var. isoviitae (Flatberg) Lönnell & Hassel
[Sphagnum fallax subsp. isoviitae
(Flatberg) M.O.Hill, Sphagnum isoviitae Flatberg]
All European records of Sphagnum magellanicum are referable to Sphagnum divinum or Sphagnum medium (Hassel et al. 2018). Sphagnum magellanicum
s.str. is confined to southern South America.
Sphagnum mirum is an Arctic species recently found in Nenets Province, European Russia (Laine et al. 2018).
166
Sphagnum beothuk is a North American species (Andrus 2006), recently found to occur in oceanic parts of Europe (Kyrkjeeide et al. 2015).
167
Shaw et al. (2012) showed that Sphagnum concinnum and Sphagnum fimbriatum are distinct species on both morphological and molecular evidence.
168
Sphagnum nitidulum is a doubtful species in need of revision, that grows around fumaroles on the island of Terceira in the Azores. It appears distinctive,
but the diagnostic characters might merely be the result of these extreme conditions (Séneca and Söderström 2009).
169
Sphagnum tescorum was found in subarctic European Russia in 2016 (Laine et al. 2018).
170
Sphagnum venustum was described from eastern Canada (Flatberg 2008). It is amphi-Atlantic, with a single location found recently in Norway (Laine et al. 2018).
171
Sphagnum viride was treated as a synonym of Sphagnum cuspidatum by Hill et al. (2006), then as a variety by Lönnell and Hassel (2018). Hanssen et al.
(2000) found that the molecular evidence for differentiation is slight. They further suggested that ”the different distribution patterns found in northwestern Europe speak in favour of a separation at the subspecies rank, but we consider a formal taxonomic decision premature at this point”. Laine
et al. (2018) state that ”the taxonomic status of this species is still somewhat unclear in relation to Sphagnum cuspidatum”. We follow Lönnell and
Hassel (2018) pending further molecular work.
172
The new combinations for Sphagnum brevifolium and Sphagnum isoviitae were proposed by Lönnell and Hassel (2018), although further molecular studies
are needed to clarify their status conclusively. It has been observed that the three varieties of Sphagnum fallax are morphologically rather stable in each of
their preferred habitats, and it may be that epigenetic processes are involved in the expression of Sphagnum fallax phenotypes.
165
JOURNAL OF BRYOLOGY
48
49
50
51
52
53
54
55
56
57
58
S. flexuosum Dozy & Molk.
S. jensenii H.Lindb.
S. lenense H.Lindb. ex L.I.Savicz
S. lindbergii Schimp.
S. majus (Russow) C.E.O.Jensen
a subsp. majus
b subsp. norvegicum Flatberg
S. obtusum Warnst.
S. pulchrum (Lindb. ex Braithw.) Warnst.
S. recurvum P.Beauv.173
S. riparium Ångstr.
S. tenellum (Brid.) Pers. ex Brid.
S. troendelagicum Flatberg
ANDREAEOPSIDA J.H.SCHAFFN.
Andreaeales Limpr.
Andreaeaceae Dumort.
2 Andreaea Hedw.
section Chasmocalyx Lindb. ex Braithw.
1 A. nivalis Hook.
section Nerviae Cardot ex G.Roth
2 A. blyttii Schimp.
3 A. crassinervia Bruch
4 A. frigida Huebener
5 A. heinemannii Hampe & Müll.Hal.
a subsp. crassifolia (Luisier) Sérgio
b subsp. heinemannii
6 A. megistospora B.M.Murray
7 A. rothii F.Weber & D.Mohr
a subsp. falcata (Schimp.) Lindb.
b subsp. rothii
section Andreaea
8 A. alpestris (Thed.) Schimp. [Andreaea rupestris
var. alpestris (Thed.) Sharp]
9 A. alpina Hedw. [Andreaea hartmanii Thed.,
Andreaea obovata Thed.]174
10 A. flexuosa R.Br. bis
a subsp. luisieri Sérgio & Sim-Sim175
11 A. hookeri Schimp. [Andreaea alpina auct.
mult.]176
12 A. mutabilis Hook.f. & Wilson
13 A. rupestris Hedw.
a var. papillosa (Lindb.) Podp.
b var. rupestris
14 A. sinuosa B.M.Murray
29
Oedipodiales Goffinet & W.R.Buck
Oedipodiaceae Schimp.
3 Oedipodium Schwägr.
1 O. griffithianum (Dicks.) Schwägr.
TETRAPHIDOPSIDA GOFFINET & W.R.BUCK
Tetraphidales M.Fleisch.
Tetraphidaceae Schimp.
4 Tetraphis Hedw.
1 T. pellucida Hedw.
5 Tetrodontium Schwägr.
1 T. brownianum (Dicks.) Schwägr.
2 T. ovatum (Funck) Schwägr.
3 T. repandum (Funck) Schwägr.
POLYTRICHOPSIDA DOWELD
Polytrichales M.Fleisch.
Polytrichaceae Schwägr.
6 Alophosia Card.
1 A. azorica (Renauld & Cardot) Cardot
7 Atrichum P.Beauv.
1 A. androgynum (Müll.Hal.) A.Jaeger177
2 A. angustatum (Brid.) Bruch & Schimp.
3 A. crispum (James) Sull.
4 A. flavisetum Mitt.
5 A. tenellum (Röhl.) Bruch & Schimp.
6 A. undulatum (Hedw.) P.Beauv.
8 Oligotrichum DC.
1 O. hercynicum (Hedw.) Lam. & DC.
9 Pogonatum P.Beauv.
1 P. aloides (Hedw.) P.Beauv.
2 P. dentatum (Menzies ex Brid.) Brid.
3 P. nanum (Hedw.) P.Beauv.
4 P. neesii (Müll.Hal.) Dozy
5 P. urnigerum (Hedw.) P.Beauv.
10
Polytrichastrum G.L.Sm.178
1 P. alpinum (Hedw.) G.L.Sm. [Polytrichastrum
norwegicum (Hedw.) Schljakov, Polytrichum alpinum Hedw., Polytrichum
alpinum var. arcticum (Sw. ex Brid.)
Wahlenb.]
2 P. altaicum Ignatov & G.L.Merr.179
OEDIPODIOPSIDA GOFFINET & W.R.BUCK
173
Sphagnum recurvum s.str. is essentially an American taxon recently found in the Azores (Dias et al. 2009). Historically the name was used in a wider sense,
encompassing Sphagnum fallax, Sphagnum angustifolium and Sphagnum flexuosum.
The confused history of Andreaea alpina and Andreaea hookeri was elucidated by Price and Ellis (2018), who describe the problems associated with the
name ‘Andreaea alpina’ and the type specimens associated with that name. In consequence, the species previously and widely known as ‘Andreaea alpina’
must now be called Andreaea hookeri, while Andreaea alpina, for which a lectotype is designated by Price and Ellis (2018), applies to the species previously
known as Andreaea obovata.
175
Andreaea flexuosa subsp. luisieri is a new taxon described from Madeira (Sérgio and Sim-Sim 2012); subsp. flexuosa occurs in the Southern Hemisphere.
176
See footnote on Andreaea alpina, above. Andreaea hookeri is therefore the correct name for the plant generally known previously as Andreaea alpina.
177
Atrichum androgynum was reported from Macaronesia and Portugal by Sérgio et al. (2010).
178
The composition of Polytrichastrum in Russia was clarified by Ivanova et al. (2014).
179
Polytrichastrum altaicum was described from the Altai Mountains in Asiatic Russia (Ignatov and Smith Merrill (1995), and later also located in European
Russia and Finnish Lapland (Ivanova et al. 2014).
174
N. G. HODGETTS ET AL.
30
3
4
5
6
11
180
P. fragile (Bryhn) Schljakov [Polytrichastrum
alpinum var. fragile (Bryhn) D.G.Long,
Polytrichum fragile Bryhn]180
P. septentrionale (Brid.) E.I.Ivanova, N.E.Bell &
Ignatov [Polytrichastrum alpinum var.
septentrionale (Sw. ex Brid.) G.L.Sm.,
Polytrichum septentrionale Brid.]181
P. sexangulare (Brid.) G.L.Sm. [Polytrichum sexangulare Hedw.]
P. sphaerothecium (Besch.) J.-P.Frahm [Polytrichastrum sexangulare var. vulcanicum
(C.E.O.Jensen) G.L.Merr., Polytrichum
sphaerothecium (Besch.) Müll.Hal.]
Polytrichum Hedw.
1 P. commune Hedw. [Polytrichum commune
var. commune, Polytrichum commune
var. uliginosum Wallr., Polytrichum uliginosum (Wallr.) Schriebl]182
2 P. densifolium Wilson ex Mitt. [Polytrichastrum formosum var. densifolium
(Wilson ex Mitt.) Z.Iwats. & Nog., Polytrichum formosum var. densifolium
(Wilson ex Mitt.) Osada]183
3 P. formosum Hedw. [Polytrichastrum formosum (Hedw.) G.L.Sm.]184
4 P. hyperboreum R.Br.
5 P. jensenii I.Hagen
6 P. juniperinum Hedw.
7 P. longisetum Sw. ex Brid. [Polytrichastrum
longisetum (Sw. ex Brid.) G.L.Sm.]185
8 P. pallidisetum Funck [Polytrichastrum pallidisetum (Funck) G.L.Sm.]186
9 P. perigoniale Michx. [Polytrichum commune
var. humile Sw., Polytrichum commune
var. perigoniale (Michx.) Hampe]187
10 P. piliferum Hedw.
11 P. strictum Menzies ex Brid. [Polytrichum
alpestre Hoppe]
12 P. swartzii Hartm. [Polytrichum commune var.
swartzii (Hartm.) Nyholm]
12
Psilopilum Brid.
1 P. cavifolium (Wilson) I.Hagen
2 P. laevigatum (Wahlenb.) Lindb.
BRYOPSIDA PAX
Buxbaumiales M.Fleisch.
Buxbaumiaceae Schimp.
13 Buxbaumia Hedw.
1 B. aphylla Hedw.
2 B. viridis (Moug. ex Lam. & DC.) Brid. ex Moug.
& Nestl.
Diphysciales M.Fleisch.
Diphysciaceae M.Fleisch.
14 Diphyscium D.Mohr
1 D. foliosum (Hedw.) D.Mohr
Timmiales Ochyra
Timmiaceae Schimp.
15 Timmia Hedw.
section Timmiaurea Brassard
1 T. austriaca Hedw.
section Timmia
2 T. bavarica Hessl.
3 T. megapolitana Hedw.
section Norvegica Brassard
4 T. comata Lindb. & Arnell
5 T. norvegica J.E.Zetterst.
6 T. sibirica Lindb. & Arnell
Encalyptales Dixon
Encalyptaceae Schimp.
16 Bryobrittonia Williams
1 B. longipes (Mitt.) D.G.Horton
Polytrichastrum fragile was treated at species level by Ivanova et al. (2014).
Polytrichastrum septentrionale was treated at species level by Ivanova et al. (2014).
Polytrichum commune var. uliginosum Wallr. was raised to specific rank by Schriebl (1991) following his observation of stability in the described diagnostic
features in culture. Hill et al. (2006) accepted this view, supported partially by Bijlsma et al. (2000), who found two distinct genetic entities within Polytrichum commune s.lat. that they recognised as Polytrichum uliginosum and Polytrichum commune, with Polytrichum commune var. perigoniale (Michx.)
Hampe treated under Polytrichum commune. Subsequent molecular and herbarium studies (Bell et al. 2010; Kariyawasam et al. in prep), have
confirmed that these two genetic entities exist, although they show that Polytrichum uliginosum should be treated under Polytrichum commune while
Polytrichum perigoniale Michx. is the correct name for the other entity. This is because Bijlsma et al. (2000) based their sampling of Polytrichum
commune on Schriebl’s concept of the species, with specimens corresponding to Polytrichum commune var. perigoniale (as can be seen in their Figure
1C), while the concept of Polytrichum uliginosum they used presumably included plants corresponding morphologically to Polytrichum commune var.
commune. The type of Polytrichum commune in the Hedwig-Schwägrichen-herbarium comprises nine stems on a single herbarium sheet. Some of
these appear to have been added by Schwägrichen for comparison purposes and are assignable to Polytrichum juniperinum, Polytrichum appressum (=Polytrichum subpilosum) and Polytrichum perigoniale. Six others match Hedwig’s original description of Polytrichum commune and descriptions in the literature
he cited, as well as the concept of Polytrichum commune var. commune traditionally used in Europe. Of these, one will be selected as a lectotype for Polytrichum commune (Kariyawasam, in prep). See also footnote to Polytrichum perigoniale, below.
183
Originally described in 1859 from Sikkim in India (Mitten 1859), Polytrichum densifolium was overlooked in Europe (and elsewhere), where specimens were
assigned to Polytrichum formosum. Most Russian specimens previously named Polytrichum formosum were transferred to Polytrichum densifolium by Ivanova
et al. (2015), and it seems likely that many other eastern European occurrences of ‘Polytrichum formosum’ may also prove to be Polytrichum densifolium.
184
Polytrichastrum formosum was returned to Polytrichum by Bell and Hyvönen (2010), a treatment followed by Ivanova et al. (2014) and others.
185
Polytrichastrum longisetum was returned to Polytrichum by Bell and Hyvönen (2010), a treatment followed by Ivanova et al. (2014) and others.
186
Polytrichastrum pallidisetum was returned to Polytrichum by Bell and Hyvönen (2010), a treatment followed by Ivanova et al. (2014) and others.
187
Molecular evidence (e.g. Bell et al. 2010; Kariyawasam et al. in prep.) clearly shows Polytrichum perigonale to be more closely related to a number of nonEuropean species than to Polytrichum commune s.str., thus necessitating its recognition as a full species rather than a variety of Polytrichum commune. The
type (G) has been examined by Kariyawasam.
181
182
JOURNAL OF BRYOLOGY
17
Encalypta Hedw.
section Streptotheca (Kindb.) Broth.
1 E. procera Bruch
2 E. streptocarpa Hedw.
section Pyromitrium Wallr. ex Hampe
3 E. alpina Sm.
4 E. mutica I.Hagen
section Rhabdotheca Müll.Hal.
5 E. pilifera Funck [Encalypta obovatifolia
Nyholm, Encalypta intermedia Jur.]188
6 E. rhaptocarpa Schwägr.
7 E. spathulata Müll.Hal. [Encalypta rhaptocarpa var. spathulata (Müll.Hal.) Husn.]
8 E. trachymitria Ripart [Encalypta rhaptocarpa
var. leptodon Lindb., Encalypta rhaptocarpa var. trachymitria (Ripart) Wijk &
Margad.]189
9 E. vulgaris Hedw.
section Megasporae D.G.Horton
10 E. longicolla Bruch
section Encalypta
11 E. affinis R.Hedw.
a subsp. affinis
b subsp. macounii (Austin) D.G.Horton
12 E. brevicolla (Bruch & Schimp.) Ångstr.
13 E. brevipes Schljakov
14 E. ciliata Hedw.
15 E. microstoma Bals.-Criv. & De Not.
Funariales M.Fleisch.
Funariaceae Schwägr.
Pyramiduloideae O.Werner, Ros & Goffinet190
18 Goniomitrium Hook.f. & Wilson
1 G. seroi Casas
19
Pyramidula Brid.
1 P. tetragona (Brid.) Brid.
Funarioideae Broth.
20 Entosthodon Schwägr.
subgenus Entosthodon
1 E. abramovae Fedosov & Ignatova191
188
31
2
E. attenuatus (Dicks.) Bryhn [Entosthodon
templetonii (Sm.) Schwägr.]
3 E. commutatus Durieu & Mont. [Entosthodon
krausei Besch.]192
4 E. dagestanicus Fedosov & Ignatova193
5 E. duriaei Mont.194
6 E. handelii (Schiffn.) Laz. [Funaria handelii
Schiffn.]195
7 E. hungaricus (Boros) Loeske
8 E. kroonkurk Dirkse & Brugués196
9 E. obtusus (Hedw.) Lindb.
10 E. stenophyllus Fedosov & Ignatova197
subgenus Plagiodus (Mitt.) Fife
11 E. convexus (Spruce) Brugués
12 E. muhlenbergii (Turner) Fife
13 E. pulchellus (H.Philib.) Brugués
14 E. schimperi Brugués
subgenus Murcia Fife
15 E. fascicularis (Hedw.) Müll.Hal.
16 E. mouretii (Corb.) Jelenc
21
Funaria Schwägr.198
1 F. aequidens Lindb. ex Broth.
2 F. arctica (Berggr.) Kindb.
3 F. hygrometrica Hedw.
4 F. microstoma Bruch ex Schimp.
22
Funariella Sérgio
1 F. curviseta (Schwägr.) Sérgio
23
Physcomitrium (Brid.) Brid. [Aphanorrhegma Sull.,
Ephemerella Mull.Hal., Physcomitrella Bruch
& Schimp., Physcomitridium G.Roth]199
1 P. arenicola Laz.
2 P. eurystomum Sendtn.
a subsp. acuminatum (Bruch & Schimp.)
Giacom.
b subsp. eurystomum
3 P. patens (Hedw.) Mitt. [Aphanorrhegma
patens (Hedw.) Lindb., Physcomitrella
patens (Hedw.) Bruch & Schimp.]
4 P. pyriforme (Hedw.) Bruch & Schimp.
Molecular studies by Fedosov (2012) showed that Encalypta pilifera is one of the most isolated taxa in sect. Rhabdotheca, thus justifying specific status, and
Encalypta pilifera is the earliest published name.
189
Fedosov (2012) proposed the recognition of Encalypta trachymitria at specific level on the basis of molecular studies on Russian material.
190
The position of Goniomitrium and Pyramidula in a subfamily within the Funariaceae was confirmed using molecular data by Werner, Ros, et al. (2007).
191
Entosthodon abramovae was described from the Caucasus by Fedosov, Ignatova, et al. (2010).
192
Entosthodon krausei was synonymised with Entosthodon commutatus by Brugués and Sérgio (2010).
193
Entosthodon dagestanicus was described from the Caucasus by Fedosov, Ignatova, et al. (2010).
194
According to Ros, Mazimpaka, et al. (2013), all occurrences of the epithet ‘durieui’, and combinations based on it, need to be changed to ‘duriaei’, because
that is the original spelling.
195
Entosthodon handelii was reported from the Caucasus new to Europe by Fedosov, Ignatova, et al. (2010).
196
Entosthodon kroonkurk was described from Spain and the Canary Islands by Dirkse and Brugués (2010).
197
Entosthodon stenophyllus was described from the Caucasus by Fedosov, Ignatova, et al. (2010).
198
Funaria anomala Jur. is a little-known taxon described from Cyprus by Juratzka in Unger and Kotschy (1865). It has also been reported from Egypt and SW
Asia. Its status is doubtful. Loeske (1929), who examined the type collection, suggested that it could be a hybrid between female Entosthodon templetonii
(Entosthodon attenuatus) and male Funaria dentata (Funaria muhlenbergii s.lat.). There have been no subsequent reports from Cyprus, and the taxon is
therefore not included in the checklist.
199
Medina et al. (2019) concluded that there are no morphological or molecular grounds for maintaining Physcomitrella and Physcomitridium as separate
genera from Physcomitrium.
32
N. G. HODGETTS ET AL.
5
6
P. readeri Müll.Hal. [Ephemerella readeri
Müll.Hal.,
Physcomitrella
readeri
(Müll.Hal.) I.G.Stone & G.A.M.Scott, Physcomitridium readeri (Müll.Hal.) G.Roth]200
P. sphaericum (C.F.Ludw. ex Schkuhr) Brid.
Disceliaceae Schimp.
24 Discelium Brid.
1 D. nudum (Dicks.) Brid.
Gigaspermales Goffinet, Wickett, O.Werner, Ros,
A.J.Shaw & C.J.Cox
Gigaspermaceae Lindb.
25 Gigaspermum Lindb.
1 G. mouretii Corb.
26
Oedipodiella Dixon
1 O. australis (Wager & Dixon) Dixon
Schwägr.) Schimp., Dicranoweisia crispula var. compacta (Schwägr.) Lindb.]
2 H. crispulum (Hedw.) Ochyra [Dicranoweisia
crispula (Hedw.) Milde]
3 H. mulahaceni (Höhn.) Ochyra [Dicranoweisia
crispula var. intermedia (J.J.Amann)
Podp.,
Dicranoweisia
intermedia
J.J.Amann, Hymenoloma intermedium
(J.J.Amann) Ochyra]205
Flexitrichaceae Ignatov & Fedosov206
31 Flexitrichum Ignatov & Fedosov
1 F. flexicaule (Schwägr.) Ignatov & Fedosov
[Ditrichum
flexicaule
(Schwägr.)
Hampe]
2 F. gracile (Mitt.) Ignatov & Fedosov [Ditrichum
crispatissimum (Müll.Hal.) Paris, Ditrichum gracile (Mitt.) Kuntze]
Catoscopiales Ignatov & Ignatova
Catoscopiaceae Broth.
27 Catoscopium Brid.
1 C. nigritum (Hedw.) Brid.
Bryoxiphiaceae Besch.
32 Bryoxiphium Brid.
1 B. madeirense Á.Löve & D.Löve
2 B. norvegicum (Brid.) Mitt.
Dicranales H.Philib ex M.Fleisch.201
Timmiellaceae Y.Inoue & H.Tsubota202
28 Timmiella (De Not.) Limpr.
1 T. anomala (Bruch & Schimp.) Limpr.
2 T. barbuloides (Brid.) Mönk.
3 T. flexiseta (Bruch) Limpr.
Archidiaceae Schimp.
33 Archidium Brid.
1 A. alternifolium (Hedw.) Mitt.
Distichiaceae Schimp.203
29 Distichium Bruch & Schimp.
1 D. capillaceum (Hedw.) Bruch & Schimp.
2 D. hagenii Ryan ex H.Philib.
3 D. inclinatum (Hedw.) Bruch & Schimp.
Hymenolomataceae Ignatov & Fedosov204
30 Hymenoloma Dusén
1 H. compactum (Schleich. ex Schwägr.) Ochyra
[Dicranoweisia compacta (Schleich. ex
200
Micromitriaceae Smyth ex Goffinet & Budke207
34 Micromitrium Austin
1 M. tenerum (Bruch & Schimp.) Crosby
Leucobryaceae Schimp.
35 Atractylocarpus Mitt.
1 A. alpinus (Schimp. ex Milde) Lindb.
2 A. subporodictyon (Broth.) Bonfim Santos &
M.Stech [Campylopus subporodictyon
(Broth.) B.H.Allen & Ireland, Dicranodontium subporodictyon Broth., Dicranum subporodictyon (Broth.) C.Gao &
T.Gao]208
Physcomitrium readeri was reported, as Ephemerella readeri, new to Europe by Hooper et al. (2010). It was moved to Physcomitridium by Goffinet and Buck
(2011), and is now in Physcomitrium (Medina et al. 2019).
201
The order Dicranales is here pragmatically broadened to include the basal lineages of haplolepidous mosses (‘Protohaplolepidae’ of Hedderson et al. 2004).
This decision necessarily results in merging the order Pottiales with Dicranales, although we retain Grimmiales as one of the crown groups of Dicranidae.
202
Inoue and Tsubota (2014, 2016) showed through molecular studies that Timmiella should be accommodated within a new family, Timmiellaceae, along
with the tropical genus Luisierella. The evidence also suggests that the family does not belong to the Pottiales but rather to the basal haplolepidous mosses
(‘Protohaplolepidae’).
203
Fedosov, Fedorova, Fedosov, et al. (2016) provided molecular evidence that Distichium cannot be placed in the Ditrichaceae, They therefore resurrected
the family Distichiaceae, which had not been used for more than a century. Further molecular evidence suggests that the Distichiaceae do not belong to
the Dicranales but rather to the basal haplolepidous mosses (‘Protohaplolepidae’).
204
Fedosov, Fedorova, Troitsky, et al. (2016) place the genus Hymenoloma in a family of its own, the Hymenolomataceae. The relationships of the species in
Hymenoloma and Dicranoweisia were elucidated by Werner, Rams, et al. (2013).
205
Hymenoloma mulahaceni is a Holarctic species whose taxonomy and distribution is described by Werner, Rams, et al. (2013).
206
The family Flexitrichaceae was described as new on the basis of molecular evidence (Fedosov, Fedorova, Fedosov, et al. 2016), which places it apart from
Ditrichaceae. As with the Distichiaceae, molecular evidence suggests that the Flexitrichaceae do not belong to the Dicranales but rather to the basal haplolepidous mosses (‘Protohaplolepidae’).
207
The Micromitriaceae is a new family described by Goffinet, Budke, et al. (2011) on molecular grounds. Its affinities with Dicranidae have not yet been
clarified but it might be close to taxa basal to Leucobryaceae (cf. Bonfim Santos and Stech 2017; Stech et al. 2012).
208
Recent molecular work shows Campylopus subporodictyon to be best placed in Atractylocarpus (Bonfim Santos and Stech 2017).
JOURNAL OF BRYOLOGY
36
37
38
Campylopus Brid.
1 C. atrovirens De Not. [Campylopus atrovirens
var. falcatus Braithw.]
2 C. brevipilus Bruch & Schimp.
3 C. cygneus (Hedw.) Brid.
4 C. flaccidus Renauld & Cardot
5 C. flexuosus (Hedw.) Brid.
6 C. fragilis (Brid.) Bruch & Schimp.
7 C. gracilis (Mitt.) A.Jaeger [Campylopus schwarzii
Schimp.]
8 C. incrassatus Müll.Hal.
9 C. introflexus (Hedw.) Brid.
10 C. oerstedianus (Müll.Hal.) Mitt.
11 C. pilifer Brid.
12 C. pyriformis (Schultz) Brid. [Campylopus
pyriformis
var.
azoricus
(Mitt.)
M.F.V.Corley]
13 C. schimperi Milde [Campylopus subulatus var.
schimperi (Milde) Husn.]209
14 C. setifolius Wilson
15 C. shawii Wilson
16 C. subulatus Schimp. ex Milde210
Dicranodontium Bruch & Schimp.
1 D. asperulum (Mitt.) Broth.
2 D. denudatum (Brid.) E.Britton [Dicranodontium
denudatum
var.
alpinum
(Schimp.) I.Hagen]
3 D. uncinatum (Harv.) A.Jaeger
Leucobryum Hampe
1 L. albidum (P.Beauv.) Lindb.211
2 L. glaucum (Hedw.) Ångstr.
3 L. juniperoideum (Brid.) Müll.Hal.
Amphidiaceae M.Stech
39 Amphidium Schimp.
1 A. lapponicum (Hedw.) Schimp.
2 A. mougeotii (Schimp.) Schimp.
3 A. curvipes (Müll.Hal.) Broth. [Amphidium tortuosum auct. eur.]
Aongstroemiaceae De Not.
40 Aongstroemia Schimp.
1 A. longipes (Sommerf.) Bruch & Schimp.
209
33
41
Dichodontium Schimp.
1 D. flavescens (Dicks.) Lindb.
2 D. pellucidum (Hedw.) Schimp.
42
Diobelonella Ochyra212
1 D. palustris (Dicks.) Ochyra [Anisothecium
palustre (Dicks.) I.Hagen, Dichodontium
palustre (Dicks.) M.Stech, Dicranella
palustris (Dicks.) Crundw.]
Dicranellaceae M.Stech
43 Dicranella (Müll.Hal.) Schimp.
1 D. campylophylla (Taylor) A.Jaeger
2 D. cerviculata (Hedw.) Schimp.
3 D. crispa (Hedw.) Schimp.
4 D. grevilleana (Brid.) Schimp.
5 D. heteromalla (Hedw.) Schimp.
6 D. howei Renauld & Cardot
7 D. humilis R.Ruthe
8 D. rufescens (Dicks.) Schimp.
9 D. schreberiana (Hedw.) Dixon
10 D. staphylina H.Whitehouse
11 D. subulata (Hedw.) Schimp.
12 D. varia (Hedw.) Schimp.
44
Microcampylopus (Müll.Hal.) M.Fleisch.
1 M. laevigatus (Thér.) Giese & J.-P.Frahm
Fissidentaceae Schimp.
45 Fissidens Hedw.
subgenus Pachyfissidens (Müll.Hal.) L.Söderstr. &
A.Hagborg
section Pachyfissidens Müll.Hal.
1 F. adianthoides Hedw.
2 F. azoricus (P.de la Varde) Bizot
3 F. dubius P.Beauv. [Fissidens cristatus Wilson
ex Mitt.]
a var. dubius
b var. mucronatus (Limpr.) Kartt., Hedenäs
& L.Söderstr.213
4 F. grandifrons Brid.
5 F. osmundoides Hedw.
6 F. polyphyllus Wilson ex Bruch & Schimp.
7 F. serrulatus Brid. [Fissidens luisieri P.de la
Varde]214
8 F. taxifolius Hedw. [Fissidens taxifolius subsp.
pallidicaulis (Mitt.) Mönk]215
Campylopus schimperi is often treated as a variety of Campylopus subulatus (e.g. Meinunger and Schröder 2007) but we follow Hill et al. (2006) in retaining
it at species level. Although they are difficult to separate in some parts of their range, Campylopus schimperi is usually very distinct in size, habit and habitat.
210
Because of inconsistencies regarding their respective habitats, there is some doubt that the taxon commonly identified as Campylopus subulatus is the
same as the type specimen; this therefore requires clarification.
211
Vanderpoorten et al. (2003) suggested that Leucobryum albidum is synonymous with Leucobryum juniperoideum. However, the two species were included
by Hill et al. (2006), and in the absence of further work are retained here. The identification of the three European species is discussed by Simmel and
Poschlod (2017).
212
Ochyra, Żarnowiec, et al. (2003) proposed a new genus, Diobelonella, for Dichodontium palustre, which, while it has molecular affinities with Dichodontium
(Stech 1999), does not sit comfortably there. It is placed alongside Dichodontium in the Aongstroemiaceae by Frey and Stech (2009).
213
Fissidens dubius var. mucronatus is morphologically characterised by the mucronate apex of the upper leaves. It also has a different ecology. Recent Dutch
research also shows differences in DNA sequences between var. mucronatus and the type (H. N. Siebel and M. Stech, pers. comm. 2019).
214
Fissidens luisieri may be distinct but further work is necessary. For the present we follow Werner, Patiño, et al. (2009) in treating it as a synonym of Fissidens
serrulatus.
215
Fissidens taxifolius subsp. pallidicaulis intergrades completely with the type subspecies and is therefore treated as a synonym.
34
N. G. HODGETTS ET AL.
section Amblyothallia (Müll.Hal.) Pursell & Brugg.Nann.
9 F. asplenioides Hedw.
subgenus Octodiceras (Brid.) Broth. [Octodiceras Brid.]
10 F. fontanus (Bach.Pyl.) Steud. [Octodiceras fontanum (Bach.Pyl.) Lindb.]
subgenus Fissidens
11 F. arcticus Bryhn
12 F. arnoldii R.Ruthe
13 F. bryoides Hedw.
a var. bryoides
b var. caespitans Schimp. [Fissidens bryoides
var. curnovii (Mitt.) J.J.Amann, Fissidens curnovii Mitt.]
14 F. coacervatus Brugg.-Nann.
15 F. crassipes Wilson ex Bruch & Schimp.
a subsp. crassipes
b subsp. warnstorfii (M.Fleisch.) Brugg.Nann.
16 F. crispus Mont. [Fissidens herzogii R.Ruthe ex
Herzog, Fissidens limbatus Sull., Fissidens minutulus Sull.]
17 F. curvatus Hornsch. [Fissidens algarvicus
Solms]
18 F. gracilifolius Brugg.-Nann. & Nyholm [Fissidens viridulus var. tenuifolius (Boulay)
A.J.E.Sm.]
19 F. gymnandrus Buse
20 F. incurvus Starke ex Röhl. [Fissidens viridulus
var. incurvus (Starke ex Röhl.) Waldh.]216
21 F. jansenii Sérgio & Pursell
22 F. monguillonii Thér.
23 F. ovatifolius R.Ruthe
24 F. pusillus (Wilson) Milde [Fissidens viridulus
var. pusillus Wilson]
25 F. rivularis (Spruce) Schimp.
26 F. rufulus Bruch & Schimp.
27 F. sublimbatus Grout
28 F. sublineaefolius (P.de la Varde) Brugg.Nann.
29 F. viridulus (Sw.) Wahlenb. [Fissidens bambergeri Schimp. ex Milde, Fissidens exiguus
auct. eur., Fissidens viridulus var. bambergeri (Schimp. ex Milde) Waldh.]217
subgenus Aloma Kindb.
30 F. celticus Paton
31 F. exilis Hedw.
216
32 F. microstictus Dixon & Luisier
33 F. nobreganus Dixon & Luisier
34 F. serratus Müll.Hal.
Dicranaceae Schimp.
Dicranoideae Lindb.
46 Dicranoloma (Renauld) Renauld
1 D. menziesii (Taylor) Broth. ex Renauld218
47
Dicranum Hedw.219
section Dicranum
1 D. bonjeanii De Not. [Dicranum undulatum
Turner, nom. illeg.]
2 D. crassifolium Sérgio, Ochyra & Séneca
3 D. leioneuron Kindb.
4 D. majus Sm.
5 D. polysetum Sw. ex anon. [Dicranum
rugosum (Hoffm. ex Funck) Brid., Dicranum undulatum Ehrh. ex F.Weber &
D.Mohr, nom. illeg.]
6 D. scoparium Hedw.
7 D. transsylvanicum Lüth
section Spuria Bruch & Schimp.
8 D. acutifolium (Lindb. & Arnell) C.E.O.Jensen
[Dicranum bergeri var. acutifolium Lindb.
& Arnell, Dicranum muehlenbeckii var.
acutifolium (Lindb. & Arnell) Nyholm]
9 D. bardunovii Tubanova & Ignatova220
10 D. brevifolium (Lindb.) Lindb. [Dicranum
muehlenbeckii var. brevifolium Lindb.,
Dicranum muehlenbeckii var. cirrhatum
(Schimp.) Lindb.]
11 D. dispersum Engelmark
12 D. drummondii Müll.Hal.
13 D. septentrionale Tubanova & Ignatova221
14 D. spurium Hedw.
15 D. undulatum Schrad. ex Brid. [Dicranum
bergeri Blandow]
section Fuscescentiformia (Kindb.) Ochyra
16 D. flexicaule Brid. [Dicranum congestum Brid.,
Dicranum fuscescens var. congestum
(Brid.) Husn., Dicranum fuscescens var.
flexicaule (Brid.) Wilson]
17 D. fuscescens Sm.
section Convolutifolia (Kindb.) Ochyra
18 D. angustum Lindb.
19 D. muehlenbeckii Bruch & Schimp.
Although morphological differences with Fissidens viridulus are small, recent molecular studies show that the closest relative of Fissidens incurvus is not
Fissidens viridulus (H. N. Siebel and M. Stech, pers. comm. 2019). It is therefore reinstated as a species.
217
The status of Fissidens bambergeri and Fissidens exiguus, treated here as synonyms of Fissidens viridulus, remains problematic. In a recent DNA study, it
appeared that an incompletely limbate species of the Fissidens bryoides complex from the Netherlands was different from Fissidens viridulus (H. N. Siebel
and M. Stech, pers. comm. 2019). However, several incompletely limbate species in this complex have been described. A morphological and molecular
revision of these species is needed.
218
Dicranoloma menziesii, a native of Australia and New Zealand, has been recorded on tree ferns in southern Ireland (Holyoak and Lockhart 2009).
219
Although this sectional classification of Dicranum is retained for now, the findings of Lang et al. (2015) suggest that it will have to be revised.
220
Dicranum bardunovii was described from Siberia (Tubanova and Ignatova 2011) but later found in the European part of the Urals (E. Ignatova pers. comm.
2018).
221
Dicranum septentrionale was described from Russia by Tubanova et al. (2010). It has since been found in Scandinavia and Austria.
JOURNAL OF BRYOLOGY
20 D. schljakovii Ignatova & Tubanova222
21 D. spadiceum J.E.Zetterst.
section Elongata I.Hagen
22 D. elongatum Schleich. ex Schwägr.
23 D. fragilifolium Lindb.
24 D. groenlandicum Brid.
25 D. laevidens R.S.Williams
section Crassinervia G.Roth
26 D. fulvum Hook. [Orthodicranum fulvum
(Hook.) G.Roth ex Casares-Gil]
27 D. scottianum Turner [Dicranum canariense
Hampe ex Müll.Hal., Orthodicranum
scottianum
(Turner) G.Roth ex
Casares-Gil]223
28 D. viride (Sull. & Lesq.) Lindb.
section Montana Hartm.
29 D. flagellare Hedw. [Orthodicranum flagellare
(Hedw.) Loeske]
30 D. montanum Hedw. [Orthodicranum montanum (Hedw.) Loeske]
31 D. tauricum Sapjegin [Dicranum strictum
Schleich. ex D.Mohr, hom. illeg., Orthodicranum tauricum (Sapjegin) Smirnova]
48
Paraleucobryum (Limpr.) Loeske
1 P. enerve (Thed.) Loeske
2 P. longifolium (Hedw.) Loeske
3 P. sauteri (Bruch & Schimp.) Loeske [Paraleucobryum longifolium var. sauteri (Bruch &
Schimp.) C.E.O.Jensen]
52
Cynodontium Bruch & Schimp.
1 C. asperifolium (Lindb. ex Arnell) Paris
2 C. bruntonii (Sm.) Bruch & Schimp.
3 C. fallax Limpr.
4 C. gracilescens (F.Weber & D.Mohr) Schimp.
5 C. jenneri (Schimp.) Stirt.
6 C. polycarpon (Hedw.) Schimp.
7 C. strumiferum (Hedw.) Lindb.
8 C. suecicum (Arnell & C.E.O.Jensen) I.Hagen
9 C. tenellum (Schimp.) Limpr.
53
Dicranoweisia Milde
1 D. cirrata (Hedw.) Lindb.
54
Glyphomitrium Brid.
1 G. daviesii (Dicks.) Brid.
55
Kiaeria I.Hagen
1 K. blyttii (Bruch & Schimp.) Broth.
2 K. falcata (Hedw.) I.Hagen
3 K. glacialis (Berggr.) I.Hagen
4 K. riparia (H.Lindb.) M.F.V.Corley [Dicranella
riparia (H.Lindb.) Mårtensson &
Nyholm, Oncophorus riparius H.Lindb.]
5 K. starkei (F.Weber & D.Mohr) I.Hagen
56
Oncophorus (Brid.) Brid.
1 O. demetrii (Renauld & Cardot) Hedenäs224
2 O. dendrophilus Hedd. & Blockeel225
3 O. elongatus (I.Hagen) Hedenäs226
4 O. integerrimus Hedenäs [Oncophorus virens
var. elongatus Limpr.]227
5 O. virens (Hedw.) Brid.
6 O. wahlenbergii Brid. [Oncophorus compactus
(Bruch & Schimp.) Kindb., Oncophorus
wahlenbergii var. compactus (Bruch &
Schimp.) Braithw.]228
57
Oreas Brid.
1 O. martiana (Hoppe & Hornsch.) Brid.
58
Oreoweisia (Bruch & Schimp.) De Not.
1 O. torquescens (Hornsch. ex Brid.) Wijk &
Margad.
59
Rhabdoweisia Bruch & Schimp.
1 R. crenulata (Mitt.) H.Jameson
2 R. crispata (Dicks.) Lindb.
3 R. fugax (Hedw.) Bruch & Schimp.
Calymperaceae Kindb.
49 Calymperes Michx.
1 C. erosum Müll.Hal.
Rhabdoweisiaceae Limpr. [Oncophoraceae M.Stech]
50
51
222
Arctoa Schimp.
1 A. anderssonii Wich.
2 A. fulvella (Dicks.) Bruch & Schimp.
3 A. hyperborea (Gunnerus ex Dicks.) Bruch &
Schimp.
Cnestrum I.Hagen
1 C. alpestre (Wahlenb. ex Huebener) Nyholm ex
Mogensen
2 C. glaucescens (Lindb. & Arnell) Holmen ex
Mogensen & Steere
3 C. schisti (F.Weber & D.Mohr) I.Hagen
35
Dicranum schljakovii was described from Russia by Ignatova, Tubanova, et al. (2015).
Price et al. (2019) clarified the correct authorship of Dicranum scottianum. Recent molecular work suggests that Dicranum canariense should be considered
a synonym of Dicranum scottianum (A. Vanderpoorten pers. comm. 2017). Morphological studies also support this conclusion (Price et al. 2019).
224
Oncophorus demetrii, a Scandinavian taxon, was recognised as a species by Hedenäs (2018), and has also been found in Arctic Russia.
225
Oncophorus dendrophilus was described by Hedderson and Blockeel (2006) and is so far known only from Crete and Cyprus.
226
The recognition of Oncophorus elongatus at species level was confirmed during a revision of Scandinavian Oncophorus by Hedenäs (2017a). It has also been
found in Russia and France.
227
Oncophorus integerrimus was recognised at species level by Hedenäs (2017a).
228
Oncophorus compactus was synonymised with Oncophorus wahlenbergii by Hedenäs (2018).
223
36
N. G. HODGETTS ET AL.
9 D. zonatum (Brid.) Kindb. [Ditrichum zonatum
var. scabrifolium Dixon]
Schistostegaceae Schimp.
60 Schistostega D.Mohr
1 S. pennata (Hedw.) F.Weber & D.Mohr
Bruchiaceae Schimp.
61 Bruchia Schwägr.
1 B. flexuosa (Schwägr.) Müll.Hal.
2 B. vogesiaca Nestl. ex Schwägr.
62
Trematodon Michx.
1 T. ambiguus (Hedw.) Hornsch.
2 T. brevicollis Hornsch.
3 T. laetevirens Hakelier & J.-P.Frahm
4 T. longicollis Michx.
5 T. perssoniorum P.Allorge & Thér. ex V.Allorge
Ditrichaceae Limpr.
63 Ceratodon Brid.
1 C. amazonum
Nieto-Lugilde,
O.Werner,
S.F.McDaniel & Ros229
2 C. conicus (Hampe) Lindb. [Ceratodon purpureus var. conicus (Hampe) Husn.]230
3 C. purpureus (Hedw.) Brid.
a subsp. purpureus
b subsp. stenocarpus (Bruch. & Schimp. ex
Müll.Hal.) Dixon231
64
Cheilothela Broth.
1 C. chloropus (Brid.) Broth.
65
Cleistocarpidium Ochyra & Bedn.-Ochyra
1 C. palustre (Bruch & Schimp.) Ochyra & Bedn.Ochyra [Pleuridium palustre (Bruch &
Schimp.) Bruch & Schimp.]
66
229
Ditrichum Timm ex Hampe
1 D. cornubicum Paton
2 D. heteromallum (Hedw.) E.Britton
3 D. lineare (Sw.) Lindb.
4 D. pallidum (Hedw.) Hampe
5 D. plumbicola Crundw.
6 D. punctulatum Mitt.
7 D. pusillum (Hedw.) Hampe
8 D. subulatum Hampe
67
Pleuridium Rabenh.
1 P. acuminatum Lindb.
2 P. subulatum (Hedw.) Rabenh.
68
Pseudephemerum (Lindb.) I.Hagen
1 P. nitidum (Hedw.) Loeske
69
Rhamphidium Mitt.
1 R. purpuratum Mitt.
70
Trichodon Schimp.
1 T. cylindricus (Hedw.) Schimp. [Ditrichum cylindricum (Hedw.) Grout]
Pottiaceae Schimp [Barbuloideae (Herzog) Hilp.]232
Merceyoideae Broth.
71 Scopelophila (Mitt.) Lindb.
1 S. cataractae (Mitt.) Broth.
2 S. ligulata (Spruce) Spruce
Pottioideae Broth.
72 Acaulon Müll.Hal.
1 A. casasianum Brugués & H.A.Crum
2 A. dertosense Casas, Sérgio, Cros & Brugués
3 A. fontiquerianum Casas & Sérgio
4 A. mediterraneum Limpr.
5 A. muticum (Hedw.) Müll.Hal.
6 A. piligerum (De Not.) Limpr.
7 A. triquetrum (Spruce) Müll.Hal.
73
Aloina Kindb.
1 A. aloides (Koch ex Schultz) Kindb.
2 A. ambigua (Bruch & Schimp.) Limpr.233
3 A. bifrons (De Not.) Delgad.
4 A. brevirostris (Hook. & Grev.) Kindb.
5 A. humilis M.T.Gallego, M.J.Cano & Ros
6 A. obliquifolia (Müll.Hal.) Broth.234
7 A. rigida (Hedw.) Limpr.
74
Barbula Hedw.235
1 B. unguiculata Hedw.
Ceratodon amazonum was described from Spain by Nieto-Lugilde et al. (2018).
Ceratodon conicus has been shown to have a hybrid origin (Nieto-Lugilde et al. 2018), with Ceratodon amazonum and Ceratodon purpureus as parent taxa,
and was considered by them a nothospecies, Ceratodon x conicus. However, it has evidently been forming self-sustaining populations for some time,
whereas Ceratodon amazonum is apparently a rare species confined to Spain, so it is here considered a true species of hybrid origin.
231
Nieto-Lugilde et al. (2018) presented molecular data suggesting that specimens corresponding to subsp. stenocarpus were not molecularly distinct from
subsp. purpureus, but a formal synonymy was not made, and the two taxa are retained for the present.
232
Inoue and Tsubota (2016) used molecular techniques to confirm that there are four clades within the Pottiaceae, corresponding to the subfamilies Trichostomoideae, Pottioideae, Merceyoideae and Streblotrichoideae. Accordingly, the subfamily Barbuloideae (Herzog) Hilp. is no longer used.
233
Aloina ambigua is doubtfully separate from Aloina aloides. The differentiating characters are quantitative and overlapping. Molecular work is needed to
elucidate further.
234
Although currently regarded as distinct, there is some evidence to suggest that Aloina obliquifolia may be conspecific with Aloina rigida (H. Siebel pers.
comm. 2017).
235
Other European species of Barbula were moved to the reinstated genera Hydrogonium and Streblotrichum and the newly described genus Gymnobarbula
by Kučera, Košnar, et al. (2013).
230
JOURNAL OF BRYOLOGY
75
Bryoerythrophyllum P.C.Chen
1 B. alpigenum (Venturi) P.C.Chen
2 B. caledonicum D.G.Long
3 B. campylocarpum (Müll.Hal.) H.A.Crum
4 B. duellii Blockeel236
5 B. ferruginascens (Stirt.) Giacom.
6 B. inaequalifolium (Taylor) R.H.Zander
7 B. recurvirostrum (Hedw.) P.C.Chen
8 B. rubrum (Jur. ex Geh.) P.C.Chen
1
2
3
4
76
77
78
Chenia R.H.Zander237
1 C. leptophylla (Müll.Hal.) R.H.Zander [Leptophascum leptophyllum (Müll.Hal.)
J.Guerra & M.J.Cano]
2 C. ruigtevleia Hedd. & R.H.Zander238
Cinclidotus P.Beauv.
1 C. aquaticus (Hedw.) Bruch & Schimp.
2 C. danubicus Schiffn. & Baumgartner
3 C. fontinaloides (Hedw.) P.Beauv.
4 C. riparius (Host ex Brid.) Arn. [Cinclidotus confertus Lüth]239
5 C. vivesii Ederra
Crossidium Jur.
1 C. aberrans Holz. & E.B.Bartram
2 C. crassinervium (De Not.) Jur.
3 C. davidai Catches.
4 C. geheebii (Broth.) Broth.
5 C. laevipilum Thér. & Trab.240
6 C. laxefilamentosum W.Frey & Kürschner241
7 C. squamiferum (Viv.) Jur.
a var. pottioideum (De Not.) Mönk.242
b var. squamiferum
5
6
7
8
9
10
11
12
13
79
80
236
Dialytrichia (Schimp.) Limpr.
1 D. mucronata (Brid.) Broth.
2 D. saxicola (Lamy) M.J.Cano [Dialytrichia fragilifolia (Bizot & J.Roux) F.Lara]243
Didymodon Hedw. [Aithobryum R.H.Zander, Exobryum R.H.Zander, Fuscobryum R.H.Zander,
14
15
37
Geheebia Schimp., Trichostomopsis Cardot,
Vinealobryum R.H.Zander]244
D. acutus (Brid.) K.Saito
D. asperifolius (Mitt.) H.A.Crum, Steere &
L.E.Anderson [Exobryum asperifolium
(Mitt.) R.H.Zander]
D. australasiae (Hook. & Grev.) R.H.Zander
[Trichostomopsis australasiae (Hook. &
Grev.) H.Rob.]
D. bistratosus Hébr. & R.B.Pierrot [Aithobryum
bistratosum (Hébr. & R.B.Pierrot)
R.H.Zander]
D. brachyphyllus (Sull.) R.H.Zander [Didymodon lamyanus (Schimp.) Thér., Vinealobryum brachyphyllum (Sull.) R.H.
Zander]245
D. cordatus Jur. [Vinealobryum cordatum
(Jur.) R.H.Zander]
D. eckeliae R.H.Zander [Vinealobryum eckeliae (R.H.Zander) R.H.Zander]246
D. fallax (Hedw.) R.H.Zander [Geheebia fallax
(Hedw.) R.H.Zander]
D. ferrugineus (Schimp. ex Besch.) M.O.Hill
[Geheebia ferruginea (Schimp. ex
Besch.) R.H.Zander]
D. giganteus (Funck) Jur. [Geheebia gigantea
(Funck) Boulay]
D. glaucus Ryan
a subsp. glaucus
b subsp. verbanus (W.E.Nicholson & Dixon)
Jan Kučera [Didymodon verbanus
(W.E. Nicholson & Dixon) Loeske]247
D. icmadophilus (Schimp. ex Müll.Hal.)
K.Saito
D. insulanus (De Not.) M.O.Hill [Vinealobryum
insulanum
(De
Not.)
R.H.Zander]
D. johansenii (R.S.Williams) H.A.Crum [Exobryum
johansenii
(R.S.Williams)
R.H.Zander]
D. luridus Hornsch. [Vinealobryum luridum
(Hornsch.) R.H.Zander]
Bryoerythrophyllum duellii was described by Blockeel et al. (2017), and is known from Greece, Crete and Cyprus.
The treatment of Hedderson and Zander (2008) is followed for Chenia.
Chenia ruigtevleia is a South African species (Hedderson and Zander 2008) recently identified from Spain (Ellis, Aleffi, et al. 2016). Further work is needed to
confirm that the Spanish and South African material are actually conspecific.
239
Recent molecular work has shown Cinclidotus confertus to be identical with C. riparius (M. Lüth pers. comm. 2017).
240
Crossidium laevipilum is doubtfully morphologically separable from Crossidium crassinervium, but molecular data (Kučera et al. in prep.) confirms its
identity.
241
Crossidium laxefilamentosum is doubtfully separable from Crossidium crassinervium.
242
Crossidium squamiferum var. pottioideum is doubtfully separable from var. squamiferum.
243
The combination Dialytrichia saxicola, which has priority over Dialytrichia fragilifolia, was made by Cano (2007).
244
The segregation of Aithobryum, Exobryum, Fuscobryum, Geheebia, Trichostomopsis and Vinealobryum, proposed by Zander (2013, 2019), is essentially
unnecessary or unsupported by molecular phylogenetic analyses.
245
Jiménez (2006) revised the European species of Didymodon and synonymised Didymodon lamyanus with Didymodon brachyphyllus.
246
Didymodon eckeliae was reported new to Europe by Puche et al. (2006). European plants are morphologically similar to American ones, but preliminary
molecular work by J. Kučera shows that they do not belong to the same lineages. The identity of the European plants therefore remains problematic.
247
Didymodon verbanus is a problematic taxon recorded in Europe from Switzerland, Austria, Italy and Germany. It is probably best recognised as a subspecies
of Didymodon glaucus, so the new combination is proposed above.
237
238
38
N. G. HODGETTS ET AL.
16
17
18
19
20
21
22
23
24
25
26
27
81
248
D. maschalogenus (Renauld & Cardot) Broth.
[Geheebia maschalogena (Ren. & Card.)
R.H.Zander].248
D. maximus (Syed & Crundw.) M.O.Hill [Geheebia maxima (Syed & Crindw.) R.H.Zander]
D. nicholsonii Culm. [Vinealobryum nicholsonii (Culm.) R.H.Zander]
D. rigidulus Hedw.
D. sinuosus (Mitt.) Delogne [Vinealobryum
sinuosum (Mitt.) R.H.Zander]
D. spadiceus (Mitt.) Limpr. [Geheebia spadicea
(Mitt.) R.H.Zander]
D. subandreaeoides (Kindb.) R.H.Zander [Fuscobryum subandreaeoides (Kindb.)
R.H.Zander]
D. tomaculosus (Blockeel) M.F.V.Corley
D. tophaceus (Brid.) Lisa [Geheebia tophacea
(Brid.) R.H.Zander]249
a subsp. erosus (J.A.Jiménez & J.Guerra) Jan
Kučera [Didymodon erosus J.A.Jiménez & J.Guerra]
b subsp. sicculus (M.J.Cano, Ros, GarcíaZam. & J.Guerra) Jan Kučera [Didymodon sicculus M.J.Cano, Ros,
García-Zam. & J.Guerra]
c subsp. tophaceus
i. var. anatinus Hamm.
ii. var. tophaceus
D. umbrosus (Müll.Hal.) R.H.Zander [Trichostomopsis umbrosa (Müll.Hal.) H.Rob.]
D. validus Limpr.250
D. vinealis (Brid.) R.H.Zander [Vinealobryum
vineale (Brid.) R.H.Zander]
Hennediella Paris
1 H. heimii (Hedw.) R.H.Zander [Desmatodon
wilczekii Meyl., Tortula rhodonia
R.H.Zander]251
a var. arctica (Lindb.) R.H.Zander252
b var. heimii
2 H. macrophylla (R.Br.bis) Paris
3 H. stanfordensis (Steere) Blockeel
82
Hilpertia R.H.Zander
1 H. velenovskyi (Schiffn.) R.H.Zander
83
Leptodontium (Müll.Hal.) Hampe
1 L. flexifolium (Dicks.) Hampe
2 L. gemmascens (Mitt.) Braithw.
3 L. proliferum Herzog253
4 L. styriacum (Jur.) Limpr.254
84
Microbryum Schimp. [Pottia Ehrh. ex Fürnr. p.p.]
1 M. curvicollum (Hedw.) R.H.Zander
2 M. davallianum (Sm.) R.H.Zander255
a var. commutatum (Limpr.) R.H.Zander
[Pottia commutata Limpr.]256
b var. conicum (Schleich. ex Schwägr.)
R.H.Zander [Pottia conica (Schleich.
ex Schwägr.) Fürnr., Pottia starckeana subsp. conica (Schleich. ex
Schwägr.) D.F.Chamb.]
c var. davallianum
3 M. floerkeanum (F.Weber & D.Mohr) Schimp.
4 M. fosbergii (E.B.Bartram) Ros, O.Werner &
Rams
5 M. longipes (J.Guerra J.J.Martínez & Ros)
R.H.Zander
6 M. rectum (With.) R.H.Zander
7 M. starckeanum (Hedw.) R.H.Zander [Pottia
starckeana (Hedw.)]
8 M. vlassovii (Laz.) R.H.Zander [Phascum vlassovii Laz., Tortula vlassovii (Laz.) Ros &
Herrnst.]257
85
Pseudocrossidium R.S.Williams258
1 P. hornschuchianum (Schultz) R.H.Zander
2 P. obtusulum (Lindb.) H.A.Crum & L.E.Anderson
3 P. replicatum (Taylor) R.H.Zander
4 P. revolutum (Brid.) R.H.Zander
Didymodon maschalogenus was reported new to Europe by Köckinger and van Melick (2007).
The treatment of Didymodon tophaceus and its infraspecific classification, including the reduction of Didymodon erosus and Didymodon sicculus to subspecies, follows Kučera, Blockeel, et al. (2018).
250
Jiménez (2006) clarified the status of Didymodon validus, which was not listed by Hill et al. (2006).
251
Tortula rhodonia was synonymised with Hennediella heimii by Cano (2008). Tortula rhodonia is a replacement name for Desmatodon wilczekii, but all possible type specimens seem to have been lost. Therefore a neotypification of Desmatodon wilczekii is necessary in order to complete the synonymisation.
252
Hennediella heimii var. arctica probably has little or no value but is retained for the present.
253
Leptodontium proliferum is a Southern Hemisphere species that was reported from a single site in England by Porley and Edwards (2010). No further
localities have been discovered, but plants with characters intermediate between this species and L. gemmascens have been found at another site in
England (Blockeel 2017). The identity of the English material of Leptodontium proliferum requires further study.
254
Leptodontium styriacum has been synonymised with Leptodontium flexifolium (Ignatov, Bersanova, et al. 2005), but it appears that Leptodontium flexifolium
has an Atlantic distribution, whereas Leptodontium styriacum has a continental one. Morphological differences between them are very small and further
work is necessary to clarify the stiuation.
255
The variability of Microbryum davallianum has not been satisfactorily clarified. Ros et al. (1996) argued for the full synonymy of the earlier recognised
varieties but no molecular data have been presented so far to support this view. In view of the mostly distinct morphologies and occasional mixed occurrences, as well as their geographical ranges, which do not fully overlap, they are retained at varietal rank pending final taxonomic clarification.
256
There are sufficient morphological differences to recognise Microbryum davallianum var. commutatum at varietal level, and the same applies to Microbryum davallianum var. conicum.
257
Microbryum vlassovii was included in Phascum by Hill et al. (2006) but placed in Tortula by Ros, Mazimpaka, et al. (2013). However, its reddish coloration
with KOH suggests that it may be more correctly placed in Microbryum.
258
Preliminary molecular studies by J. Kučera suggest that what has been recognised as Pseudocrossidium obtusulum and also the European records of Pseudocrossidium replicatum may prove to be Pseudocrossidium hornschuchianum, but until a revision is completed both species are retained.
249
JOURNAL OF BRYOLOGY
86
Pterygoneurum Jur.
1 P. kozlovii Laz.
2 P. lamellatum (Lindb.) Jur.
3 P. ovatum (Hedw.) Dixon [Pterygoneurum compactum M.J.Cano, J.Guerra & Ros,
Pterygoneurum crossidioides W.Frey,
Herrnst. & Kürschner, Pterygoneurum
squamosum Segarra & Kürschner]259
4 P. papillosum Oesau
5 P. sampaianum (Guim.) Guim.
6 P. subsessile (Brid.) Jur.
87
Stegonia Venturi
1 S. latifolia (Schwägr.) Venturi ex Broth.
a var. latifolia
b var. pilifera (Brid.) Broth.260
88
259
Syntrichia Brid.
1 S. bogotensis (Hampe) R.H.Zander
2 S. calcicola J.J.Amann
3 S. caninervis Mitt.
a var. abranchesii (Luisier) R.H.Zander
b var. astrakhanica Ignatov, Ignatova &
Suragina
c var. caninervis
d var. gypsophila (J.J.Amann ex G.Roth)
Ochyra
4 S. echinata (Schiffn.) Herrnst. & Ben-Sasson
5 S. fragilis (Taylor) Ochyra
6 S. handelii (Schiffn.) S.Agnew & Vondr.
7 S. laevipila Brid. [Syntrichia pagorum (Milde)
J.J.Amann]261
8 S. latifolia (Bruch ex Hartm.) Huebener
9 S. minor (Bizot) M.T.Gallego, J.Guerra, M.J.Cano,
Ros & Sánchez-Moya
10 S. montana Nees [Syntrichia intermedia Brid.]
a var. calva (Durieu & Sagot ex Bruch &
Schimp.) J.J.Amann
b var. montana
11 S. norvegica F.Weber
12 S. papillosa (Wilson) Jur.
13 S. papillosissima (Copp.) Loeske
14 S. princeps (De Not.) Mitt.
15 S. rigescens (Broth. & Geh.) Ochyra
39
16 S. ruraliformis (Besch.) Mans. [Syntrichia ruralis
var. ruraliformis (Besch.) Delogne]262
17 S. ruralis (Hedw.) F.Weber & D.Mohr [Syntrichia densa (Velen.) J.-P.Frahm, Syntrichia
glabra J.-P.Frahm & M.T.Gallego]263
a var. epilosa (Venturi) J.J.Amann264
b var. ruralis
18 S. sinensis (Müll.Hal.) Ochyra
19 S. submontana (Broth.) Ochyra265
20 S. subpapillosissima (Bizot & R.B.Pierrot ex
W.A.Kramer)
M.T.Gallego
&
J.Guerra266
21 S. virescens (De Not.) Ochyra
89
Tortula Hedw. [Desmatodon Brid., Phascum Hedw.,
Pottia Ehrh. ex Fürnr. p.p., Protobryum
(Dicks.) J.Guerra & M.J.Cano]
1 T. acaulon (With.) R.H.Zander [Phascum cuspidatum Hedw.]267
a var. acaulon [Phascum cuspidatum var.
cuspidatum]
b var. papillosa (Lindb.) R.H.Zander [Phascum
cuspidatum var. papillosum (Lindb.)
G.Roth]
c var. pilifera (Hedw.) R.H.Zander [Phascum
cuspidatum var. piliferum (Hedw.)
Hook. & Taylor]
d var. retortifolia (J.Guerra & Ros) R.H.Zander
[Phascum cuspidatum var. retortifolium J.Guerra & Ros]
e var. schreberiana (Dicks.) R.H.Zander
[Phascum cuspidatum var. schreberianum (Dicks.) Brid.]
2 T. amplexa (Lesq.) Steere [Syntrichia amplexa
(Lesq.) R.H.Zander]
3 T. ampliretis Crundw. & D.G.Long
4 T. atrovirens (Sm.) Lindb.
5 T. bogosica (Müll.Hal.) R.H.Zander [Desmatodon bogosicus Müll.Hal.]
6 T. bolanderi (Lesq. & James) M.Howe
7 T. brevissima Schiffn.
8 T. canescens Mont.
9 T. caucasica Broth. [Pottia intermedia (Turner)
Fürnr., Tortula modica R.H.Zander]268
Synonymies of Pterygoneurum ovatum follow Guerra, Brugués, et al. (2006). However, this treatment is not yet supported by molecular data.
Stegonia latifolia var. pilifera is reinstated because it appears to be a distinctive and stable taxon, albeit differing from the type in only a single character,
the excurrent costa.
261
Syntrichia pagorum (Milde) J.J.Amann, listed as a synonym of Syntrichia laevipila by Hill et al. (2006), may prove to be a good species, but more work is
needed (Afonina et al. 2014).
262
Molecular and morphological evidence suggest that Syntrichia ruraliformis should be regarded as a species (Hedenäs, Heinrichs, et al. 2019).
263
According to Frahm and Sabovljević (2006), Syntrichia glabra is a juvenile form of Syntrichia densa (Velen.) J.-P.Frahm. Syntrichia densa was synonymised
with Syntrichia calcicola by Gallego et al. (2002), but Kramer (1980) synonymised Tortula densa with Tortula ruralis. The latter synonymy is followed here.
264
Syntrichia ruralis var. epilosa was resurrected by Gallego et al. (2018). It may represent a special phenotype within Syntrichia ruraliformis or Syntrichia ruralis
but more study is required (Hedenäs et al. 2019).
265
Syntrichia submontana is essentially a central Asian species recorded from the Caucasus (Afonina et al. 2014).
266
Syntrichia subpapillosissima may represent a special phenotype within Syntrichia ruraliformis or Syntrichia ruralis but more study is required (Hedenäs,
Heinrichs, et al. 2019).
267
We follow the treatment of Ros, Mazimpaka, et al. (2013) and refer the species included in Phascum by Hill et al. (2006) to Tortula.
268
Tortula modica was synonymised with Tortula caucasica by Ros, Muñoz, et al. (2008).
260
N. G. HODGETTS ET AL.
40
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
269
cernua (Huebener) Lindb.
cuneifolia (Dicks.) Turner
freibergii Dixon & Loeske
guepinii (Bruch & Schimp.) Broth. [Desmatodon guepinii Bruch & Schimp.]
T. hoppeana (Schultz) Ochyra [Desmatodon
latifolius (Hedw.) Brid.]
T. inermis (Brid.) Mont.
T. israelis Bizot & F.Bilewsky [Tortula baetica
(Casas & R.Oliva) J.Guerra & Ros]
T. laureri (Schultz) Lindb. [Desmatodon laureri
(Schultz) Bruch & Schimp.]
T. leucostoma (R.Br.) Hook. & Grev. [Desmatodon leucostoma (R.Br.) Berggr.]
T. lindbergii Broth. [Pottia lanceolata (Hedw.)
Müll.Hal., Tortula lanceola R.H.Zander]269
T. lingulata Lindb.
T. marginata (Bruch & Schimp.) Spruce [Desmatodon meridionalis Luisier]
T. mucronifolia Schwägr.
T. muralis Hedw.
a subsp. muralis270
i. var. aestiva Hedw.
ii. var. muralis
b subsp. obtusifolia (Schwägr.) Culm.
[Tortula obtusifolia (Schwägr.)
Mathieu]271
T. pallida (Lindb.) R.H.Zander
T. protobryoides R.H.Zander [Protobryum
bryoides (Dicks.) J.Guerra & M.J.Cano]272
T. randii (Kenn.) R.H.Zander [Desmatodon
randii (Kenn.) Laz.)]
T. revolvens (Schimp.) G.Roth
T. schimperi M.J.Cano, O.Werner & J.Guerra
[Tortula subulata var. angustata
(Schimp.) Limpr.]
T. solmsii (Schimp.) Limpr.
T. subulata Hedw. [Tortula subulata var.
graeffii Warnst., Tortula subulata var.
subinermis (Bruch & Schimp.) Wilson]
T. systylia (Schimp.) Lindb. [Desmatodon systylius Schimp.]
T. truncata (Hedw.) Mitt.
33 T. ucrainica (Laz.) R.H.Zander [Desmatodon
ucrainicus Laz.]
34 T. vahliana (Schultz) Mont.
35 T. viridifolia (Mitt.) Blockeel & A.J.E.Sm. [Pottia
crinita Bruch & Schimp.]
36 T. wilsonii (Hook.) R.H.Zander
T.
T.
T.
T.
90
Triquetrella Müll.Hal.
1 T. arapilensis Luisier
Streblotrichoideae Y.Inoue & H.Tsubota273
91 Streblotrichum P.Beauv.274
1 S. convolutum (Hedw.) P.Beauv. [Barbula convoluta Hedw.]
a var. commutatum (Jur.) J.J.Amann [Barbula
convoluta var. sardoa Schimp., Streblotrichum commutatum (Jur.) Hilp.]275
b var. convolutum
2 S. enderesii (Garov.) Loeske [Barbula enderesii
Garov.]
Trichostomoideae Broth.
92 Anoectangium Schwägr.
1 A. aestivum (Hedw.) Mitt.
2 A. angustifolium Mitt.276
3 A. handelii Schiffn.
93
Aschisma Lindb.
1 A. carniolicum (F.Weber & D.Mohr) Lindb.
2 A. cuynetii (Bizot & R.B.Pierrot) J.Guerra &
M.J.Cano
94
Chionoloma Dixon [Paraleptodontium D.G.Long,
Oxystegus (Limpr.) Hilp.]277
1 C. bombayense (Müll.Hal.) P.Sollman [Pseudosymblepharis bombayensis (Müll.Hal.)
P.Sollman]278
2 C. cylindrotheca (Mitt.) M.Alonso, M.J.Cano &
J.A.Jiménez [Chinoloma daldinianum
(De Not.) M.Alonso, M.J.Cano & J.A.
Jiménez, Oxystegus daldinianus (De
Not.) Köckinger, O.Werner & Ros]279
3 C. hibernicum (Mitt.) M.Alonso, M.J.Cano & J.A.Jiménez [Oxystegus hibernicus (Mitt.) Hilp.,
Trichostomum hibernicum (Mitt.) Dixon]
Tortula lanceola was synonymised with Tortula lindbergii by Ros, Muñoz, et al. (2008).
The two varieties of Tortula muralis subsp. muralis intergrade but there is a certain level of distinctness, so this treatment follows Košnar and Kolář (2009).
Košnar and Kolář (2009) reduced Tortula obtusifolia to a subspecies of Tortula muralis.
272
We follow the treatment of Ros, Mazimpaka, et al. (2013) for Tortula protobryoides.
273
The subfamily Streblotrichoideae was proposed by Inoue and Tsubota (2016).
274
Streblotrichum was re-established on molecular evidence by Kučera, Košnar, et al. (2013).
275
Streblotrichum convolutum var. commutatum is retained at varietal rank. Although it is not always morphologically distinct, molecular evidence clearly
separates the two taxa (Kučera, Košnar, et al. 2013).
276
Unpublished molecular work by J. Kučera shows Anoectangium angustifolium to be distinct from Anoectangium aestivum.
277
While Oxystegus is a molecularly well-delimited group, retaining it leaves problems, notably in the delimitation of Pseudosymblepharis Broth. Therefore, we
follow Alonso et al. (2016) in placing these species in Chionoloma.
278
Chionoloma bombayense, recorded in Europe from the Caucasus, is retained in the checklist for the present, following Ignatova, Kuznetsova, et al. (2012).
However, Alonso et al. (2018) circumscribe Chionoloma bombayense in a much narrower sense, leaving many other plants in Chionoloma bombayense s.lat,
including the Caucasus plant, without a name.
279
Originally described as Didymodon cylindricus var. daldinianus De Not., Chionoloma daldinianum was treated in various ways historically, including being
omitted altogether by Hill et al. (2006); it was resurrected as a species with molecular support by Köckinger, Werner, et al. (2010), then brought into Chionoloma by Alonso et al. (2016). Later, Alonso et al. (2019) synonymised Chionoloma daldinianum with the more widespread Chionoloma cylindrotheca.
270
271
JOURNAL OF BRYOLOGY
4
5
6
95
96
280
C. minus (Köckinger, O.Werner & Ros) M.Alonso,
M.J.Cano & J.A.Jiménez [Oxystegus
minor Köckinger, O.Werner & Ros]280
C. recurvifolium (Taylor) M.Alonso, M.J.Cano &
J.A.Jiménez [Oxystegus recurvifolius
(Taylor) R.H.Zander, Paraleptodontium
recurvifolium (Taylor) D.G.Long, Trichostomum recurvifolium (Taylor)
R.H.Zander]
C. tenuirostre (Hook. & Taylor) M.Alonso,
M.J.Cano & J.A.Jiménez [Oxystegus
tenuirostris (Hook. & Taylor) A.J.E.Sm.,
Trichostomum tenuirostre (Hook. &
Taylor) Lindb.]
a var. holtii (Braithw.) M.Alonso, M.J.Cano &
J.A.Jiménez [Oxystegus tenuirostris
var. holtii (Braithw.) A.J.E.Sm.]
b var. tenuirostre
Ephemerum Hampe281
1 E. cohaerens (Hedw.) Hampe
2 E. crassinervium (Schwägr.) Hampe282
a subsp. rutheanum (Schimp.) Holyoak
[Ephemerum hibernicum Holyoak &
V.S.Bryan, Ephemerum rutheanum
Schimp., Ephemerum serratum var.
rutheanum (Schimp.) Jur.]
b subsp. sessile (Bruch) Holyoak [Ephemerum
sessile (Bruch) Müll.Hal.]
3 E. recurvifolium (Dicks.) Boulay
4 E. serratum (Hedw.) Hampe [Ephemerum
minutissimum Lindb., Ephemerum serratum var. minutissimum (Lindb.) Grout]283
5 E. spinulosum Bruch & Schimp. ex Schimp.
6 E. stoloniferum (Hedw.) L.T.Ellis & M.J.Price
[Ephemerum serratum auct., Ephemerum stellatum H.Philib.]284
Eucladium Bruch & Schimp.
41
1 E. verticillatum (With.) Bruch & Schimp. [Eucladium verticillatum var. angustifolium
Lindb.]285
97
Gymnobarbula Jan Kučera286
1 G. bicolor (Bruch & Schimp.) Jan Kučera
[Barbula bicolor (Bruch & Schimp.)
Lindb.]
98
Gymnostomum Nees & Hornsch.
1 G. aeruginosum Sm.
a var. aeruginosum
b var. obscurum J.Guerra
2 G. calcareum Nees & Hornsch. [Gymnostomum
calcareum var. lanceolatum Sérgio,
Gymnostomum lanceolatum M.J.Cano,
Ros & J.Guerra]287
a var. atlanticum Sérgio288
b var. calcareum
3 G. viridulum Brid.
99
Gyroweisia Schimp.
1 G. reflexa (Brid.) Schimp.
2 G. tenuis (Hedw.) Schimp.
100
Hydrogonium (Müll.Hal.) A.Jaeger289
1 H. amplexifolium (Mitt.) P.C.Chen [Barbula
amplexifolia (Mitt.) A.Jaeger]290
2 H. bolleanum (Müll.Hal.) A.Jaeger [Barbula
bolleana (Müll.Hal.) Broth.]
3 H. consanguineum (Thwaites & Mitt.) Hilp.
[Barbula consanguinea (Thwaites &
Mitt.) A.Jaeger]291
a var. kurilense (Ignatova & Ignatov) Jan
Kučera292
4 H. croceum (Brid.) Jan Kučera [Barbula crocea
(Brid.) F.Weber & D.Mohr]
Oxystegus minor was described as a new species by Köckinger, Werner, et al. (2010), then brought into Chionoloma by Alonso et al. (2016). However,
significant doubts remain about its distinctiveness as a species: in north-western Europe it intergrades morphologically with Chionoloma hibernicum.
All recent phylogenies (e.g. Werner, Ros, et al. 2005) agree in placing Ephemerum within the Trichostomoideae rather than its own family, however strange
this may seem morphologically.
282
The treatment of Ephemerum crassinervium and related taxa follows Holyoak (2010).
283
As a consequence of lectotypification, Ephermerum minutissimum Lindb. was placed in synonymy with Ephemerum serratum (Hedw.) Hampe (Ellis and
Price 2015)
284
Ephemerum stellatum was synonymised with Ephemerum serratum by Holyoak (2010). Subsequently, Ellis and Price (2015) lectotypified E. stoloniferum and
showed that it was the correct name for the large-spored taxon previously treated under the name E. serratum.
285
Eucladium verticillatum var. angustifolium is regarded as nothing more than a form of the type.
286
Gymnobarbula was established as a new genus by Kučera, Košnar, et al. (2013).
287
Gymnostomum lanceolatum was reduced to a variety of Gymnostomum calcareum by Sérgio (2006). We treat it as a synonym, as it appears to be merely a
form of that species.
288
Gymnostomum calcareum var. atlanticum was described by Sérgio (2006). On molecular evidence, it is a well-marked variety that may be sufficiently distinct to be elevated to specific status.
289
Hydrogonium was re-established on molecular evidence by Kučera, Košnar, et al. (2013).
290
Barbula amplexifolia was reported new to Europe by Köckinger and Kučera (2007) and transferred to Hydrogonium by Kučera, Košnar, et al. (2013).
291
Hydrogonium consanguineum was earlier reported from Europe as Barbula indica; Köckinger, Kučera, et al. (2012) proved that these reports belong to
Barbula consanguinea, transferred to Hydrogonium by Kučera, Košnar, et al. (2013).
292
The European expression of Hydrogonium consanguineum is var. kurilense; var. consanguineum occurs in Asia; var. cancellatum (Müll.Hal.) Jan Kučera is a
North American plant (Kučera, Košnar, et al. 2013).
281
42
101
N. G. HODGETTS ET AL.
Hymenostylium Brid. [Ardeuma R.H.Zander &
Hedd.]293
1 H. gracillimum (Nees & Hornsch.) Köckinger
& Jan Kučera [Gymnostomum
boreale Nyholm & Hedenäs]294
2 H. recurvirostrum (Hedw.) Dixon [Gymnostomum recurvirostrum Hedw.]
a var. insigne (Dixon) E.B.Bartram [Ardeuma
annotinum (Mitt. ex Dixon) R.H.
Zander & Brinda, Ardeuma insigne
(Dixon) R.H.Zander & Hedd., Hymenostylium insigne (Dixon) Podp.]
b var. recurvirostrum [Ardeuma recurvirostrum (Hedw.) R.H.Zander & Hedd.]
3 H. xerophilum Köckinger & Jan Kučera295
102
Hyophila Brid.
1 H. involuta (Hook.) A.Jaeger
103
Leptobarbula Schimp.
1 L. berica (De Not.) Schimp.
104
Molendoa (Müll.Hal.) Hampe
1 M. hornschuchiana (Hook.) Lindb. ex Limpr.
[Anoectangium
hornschuchianum
(Hook.) Funck ex Hornsch., Anoectangium sendtnerianum Bruch & Schimp.,
Molendoa sendtneriana (Bruch &
Schimp.) Limpr., Molendoa tenuinervis
Limpr.]296
2 M. schliephackei (Schlieph.) R.H.Zander
[Anoectangium
schliephackei
(Schlieph.) Paris]
3 M. taeniatifolia Herzog [Anoectangium taeniatifolium (Herzog) M.O.Hill]
4 M. warburgii (Crundw. & M.O.Hill) R.H.Zander
[Anoectangium warburgii (Limpr.) Paris]
105
Pottiopsis Blockeel & A.J.E.Sm.
1 P. caespitosa (Brid.) Blockeel & A.J.E.Sm. [Trichostomum pallidisetum H.Müll., Trichostomum triumphans De Not.,
293
Weissia triumphans (De Not.) M.O.Hill,
Weissia tyrrhena M.Fleisch.]297
106
Splachnobryum Müll.Hal.
1 S. obtusum (Brid.) Müll.Hal.
107
Tortella (Müll.Hal.) Limpr. [Pleurochaete Lindb.]
1 T. alpicola Dixon
2 T. x cuspidatissima (Cardot & Thér.)
O.Werner, Köckinger & Ros 298
3 T. densa (Lorentz & Molendo) Crundw. &
Nyholm [Tortella inclinata var. densa
(Lorentz & Molendo) Limpr.]
4 T. fasciculata (Culm.) Culm. [Tortella bambergeri auct., non (Schimp.) Broth.
p.p., Tortella tortuosa subsp. fasciculata Culm.]299
5 T. flavovirens (Bruch) Broth. [Tortella limosella
(Stirt.)
P.W.Richards
&
E.C.Wallace]300
a var. flavovirens
b var. glareicola (T.A.Chr.) Crundw. &
Nyholm
c var. papillosissima Sérgio & Casas
6 T. fragilis (Drumm.) Limpr.301
7 T. humilis (Hedw.) Jenn.
8 T. inclinata (R.Hedw.) Limpr.
9 T. inflexa (Bruch) Broth.
10 T. limbata (Schiffn.) Geh. & Herzog
11 T. mediterranea Köckinger, Lüth, O.Werner
& Ros302
12 T. nitida (Lindb.) Broth.
13 T. pseudofragilis (Thér.) Köckinger & Hedenäs
[Tortella bambergeri auct., non
(Schimp.) Broth. p.p., Tortella fragilis
var. moravica Podp.]303
14 T. rigens Alberts.
15 T. spitsbergensis (Bizot & Thér.) O.Werner,
Köckinger & Ros [Trichostomum arcticum Kaal.]304
16 T. squarrosa (Brid.) Limpr. [Pleurochaete
squarrosa (Brid.) Lindb.]305
The transfer of most Hymenostylium species, including Hymenostylium recurvirostrum and its varieties, to Ardeuma (Zander and Hedderson 2016) is not
supported according to molecular evidence (Kučera in prep.).
294
The position of Hymenostylium gracillimum and the synonymy of Gymnostomum boreale were clarified by Köckinger and Kučera (2011).
295
Hymenostylium xerophilum was described from Austria as a new species by Köckinger and Kučera (2011). It has since been found in Germany and Crimea.
296
Geissler (1985) proposed the synonymy of Molendoa sendtneriana and Molendoa tenuinervis with Molendoa hornschuchiana. This is followed here, as it is
supported by preliminary molecular results by J. Kučera.
297
The position of Pottiopsis caespitosa and its synonymies were clarified by Ros and Werner (2007). However, Trichostomum/Weissia triumphans is a distinctive form, and continues to be recognised by many bryologists.
298
Werner, Köckinger, et al. (2014) used molecular techniques to show that Tortella x cuspidatissima was of hybrid origin, with Tortella arctica (Arnell) Crundw.
& Nyholm and Tortella spitsbergensis as parents. Tortella arctica itself does not occur in Europe. The record of Tortella arctica from Jämtland in Sweden (Ellis,
Akhoondi Darzikolaei, et al. 2011) proved, according to the molecular data, to be a form of Tortella tortuosa (Köckinger and Hedenäs 2017).
299
Tortella fasciculata was resurrected by Köckinger and Hedenäs (2017) as part of their revision of Tortella bambergeri.
300
Further studies on Tortella limosella are necessary to determine the final position of Tortella limosella within Tortella flavovirens.
301
The authorship of Tortella fragilis was discussed in Ochyra, Lewis Smith, et al. (2008). The name should correctly be ascribed to Drummond.
302
Tortella mediterranea was described from southern Europe by Köckinger, Lüth, et al. (2018).
303
The name Tortella pseudofragilis was introduced by Köckinger and Hedenäs (2017) as part of their revision of Tortella bambergeri.
304
The identity and relationships of Tortella spitsbergensis were clarified by Werner, Köckinger, et al. (2014) using molecular techniques.
305
We follow Ros, Mazimpaka, et al. (2013) in accepting Werner, Ros, et al. (2005) and Grundmann et al. (2006), who include Pleurochaete squarrosa within the
genus Tortella.
JOURNAL OF BRYOLOGY
17
T. tortuosa (Hedw.) Limpr. [Tortella bambergeri (Schimp.) Broth.]306
108
Trichostomum Bruch.
1 T. brachydontium Bruch
2 T. crispulum Bruch
109
Weissia Hedw.
1 W. angustifolia (Baumgartner) D.A.Callaghan [Weissia longifolia var. angustifolia (Baumgartner) Crundw. &
Nyholm]307
2 W. brachycarpa (Nees & Hornsch.) Jur.
3 W. condensa (Voit) Lindb.
a var. armata (Thér. & Trab.) M.J.Cano, Ros
& J.Guerra
b var. condensa
4 W. controversa Hedw.
a var. controversa
b var. crispata (Nees & Hornsch.) Nyholm
c var. densifolia (Bruch & Schimp.) Wilson308
5 W. levieri (Limpr.) Kindb.
6 W. longifolia Mitt.
7 W. x mittenii (Bruch & Schimp.) Mitt. emend.
A.J.E.Sm.309
8 W. perssonii Kindb.
9 W. rostellata (Brid.) Lindb.
10 W. rutilans (Hedw.) Lindb.
11 W. squarrosa (Nees & Hornsch.) Müll.Hal.
12 W. sterilis W.E.Nicholson
13 W. wilsonii D.A.Callaghan [Weissia multicapsularis auct.]310
14 W. wimmeriana (Sendtn.) Bruch & Schimp.
Grimmiales M.Fleisch.
Saelaniaceae Ignatov & Fedosov311
110 Saelania Lindb.
1 S. glaucescens (Hedw.) Broth.
Seligeriaceae Schimp.
111 Blindia Bruch & Schimp.
1 B. acuta (Hedw.) Bruch & Schimp.
2 B. caespiticia (F.Weber & D.Mohr) Müll.Hal.
306
43
112
Blindiadelphus (Lindb.) Fedosov & Ignatov [Seligeria Bruch & Schimp.
subgenus Blindiadelphus Lindb.]312
1 B. campylopodus (Kindb.) Fedosov & Ignatov
[Seligeria campylopoda Kindb.]
2 B. diversifolius (Lindb.) Fedosov & Ignatov
[Seligeria diversifolia Lindb.]
3 B. polaris (Berggr.) Fedosov & Ignatov [Seligeria polaris Berggr.]
4 B. recurvatus (Hedw.) Fedosov & Ignatov [Seligeria recurvata (Hedw.) Bruch & Schimp.]
5 B. subimmersus (Lindb.) Fedosov & Ignatov
[Seligeria subimmersa Lindb.]
113
Seligeria Bruch & Schimp.
subgenus Seligeria
1 S. acutifolia Lindb.
2 S. brevifolia (Lindb.) Lindb.
3 S. calcarea (Hedw.) Bruch & Schimp.
4 S. calycina Mitt. ex Lindb. [Seligeria paucifolia auct. non (With.) Carruth.]
5 S. donniana (Sm.) Müll.Hal. [Seligeria galinae
Mogensen & I.Goldberg]313
6 S. pusilla (Hedw.) Bruch & Schimp.
subgenus Megalosporia Vitt [Trochobryum Breidl.
& Beck]
7 S. austriaca T.Schauer
8 S. carniolica (Breidl. & Beck) Nyholm [Trochobryum carniolicum Breidl. & Beck]
9 S. irrigata (H.K.G.Paul) Ochyra & Gos
10 S. oelandica C.E.O.Jensen & Medelius
11 S. patula (Lindb.) I.Hagen [Seligeria alpestris
T.Schauer, Seligeria patula var. alpestris
(T.Schauer) Gos & Ochyra, Seligeria tristichoides var. patula (Lindb.) Broth.]
12 S. trifaria (Brid.) Lindb.
a var. longifolia (Lindb. ex Broth.) Ochyra &
Gos314
b var. trifaria
13 S. tristichoides Kindb.
Ptychomitriaceae Schimp.
114 Brachydontium Fürnr.315
Using morphological and molecular methods, Köckinger and Hedenäs (2017) demonstrated that the type of Tortella bambergeri was synonymous with
Tortella tortuosa. Plants recently treated as Tortella bambergeri are referable to Tortella fasciculata and Tortella pseudofragilis. Up to now, the acceptance of
Tortella tortuosa var. fragilifolia (Jur.) Limpr. has been based on the treatment of Eckel (1998), but this concept is based on plants later referred to Tortella
bambergeri (Eckel 2010). In general, the name ‘var. fragilifolia’ has been used for morphs of Tortella tortuosa with fragile leaves, and the variety is therefore
not included in the checklist.
307
Weissia longifolia var. angustifolia was raised to species level by Callaghan et al. (2019) on the basis of morphological and molecular evidence.
308
Weissia controversa var. densifolia is retained as a variety for now, but it is very doubtfully distinct from var. controversa, and significantly less so than var. crispata.
309
Smith (2004) was the first to propose that Weissia x mittenii was a hybrid, and this was accepted by Hill et al. (2006). The name was later typified by
Callaghan (2019).
310
Weissia wilsonii was newly described by Callaghan et al. (2019) to replace the taxon which was named Weissia multicapsularis (Sm.) Mitt. by modern
authors. The type of the latter refers to Tortula acaulon, into which plants of Weissia longifolia were admixed.
311
Molecular studies by Fedosov, Fedorova, Fedosov, et al. (2016) showed that Saelania is best placed in its own family within the Grimmiales.
312
Blindiadelphus was segregated from Seligeria on the basis of morphological and molecular evidence (Fedosov, Fedorova, Ignatova, et al. 2017).
313
Seligeria galinae was synonymised with Seligeria donniana by Fedosov, Fedorova, Ignatova, et al. (2017).
314
Seligeria trifaria var. longifolia was not mentioned in Hill et al. (2006) but is regarded as a distinct taxon in the central European mountains (Ochyra and Gos 1992).
315
Molecular evidence places Brachydontium in the Ptychomitriaceae (Fedosov et al. 2017a).
44
N. G. HODGETTS ET AL.
1
B. trichodes (F.Weber) Milde [Seligeria transylvanica Plam.]
115
Campylostelium Bruch & Schimp.
1 C. pitardii (Corb.) E.Maier [Grimmia pitardii Corb.]
2 C. saxicola (F.Weber & D.Mohr) Bruch & Schimp.
3 C. strictum Solms
116
Indusiella Broth. & Müll.Hall.
1 I. thianschanica Broth. & Müll.Hal.
117
Jaffueliobryum Thér.
1 J. latifolium Lindb. & Arnell ex Thér.
118
Ptychomitrium Fürnr.
1 P. incurvum (Schwägr.) Spruce
2 P. nigrescens (Kunze) Wijk & Margad.
3 P. polyphyllum (Dicks. ex Sw.) Bruch & Schimp.
Grimmiaceae Arn.
Racomitrioideae Ochyra & Bedn.-Ochyra
119 Racomitrium Brid.316
subgenus Racomitrium
1 R. lanuginosum (Hedw.) Brid.
subgenus Niphotrichum Bedn.-Ochyra [Niphotrichum (Bedn.-Ochyra) Bedn.-Ochyra & Ochyra]
2 R. canescens (Hedw.) Brid. [Niphotrichum canescens (Hedw.) Bedn.-Ochyra & Ochyra]
a subsp. canescens
b subsp. latifolium (C.E.O.Jensen) Frisvoll
[Niphotrichum canescens subsp. latifolium (Frisvoll) Bedn.-Ochyra & Ochyra]
3 R. elongatum Ehrh. ex Frisvoll [Niphotrichum
elongatum (Ehrh. ex Frisvoll) Bedn.Ochyra & Ochyra]
4 R. ericoides (Brid.) Brid. [Niphotrichum ericoides (Brid.) Bedn.-Ochyra & Ochyra]
5 R. panschii (Müll.Hal.) Kindb. [Niphotrichumpanschii (Müll.Hal.) Bedn.-Ochyra &
Ochyra]
subgenus Cataracta Vilh. [Codriophorus P.Beauv.,
Dilutineuron Bedn.-Ochyra, Sawicki, Ochyra, Szczecińska & Plášek]
6 R. aciculare (Hedw.) Brid. [Codriophorus acicularis (Hedw.) P.Beauv.]
7 R. aquaticum (Brid. ex Schrad.) Brid. [Codriophorus aquaticus (Brid. ex Schrad.)
Bedn.-Ochyra & Ochyra]
8 R. fasciculare (Hedw.) Brid. [Codriophorus
fascicularis (Hedw.) Bedn.-Ochyra &
Ochyra,
Dilutineuron
fasciculare
316
(Hedw.)
Bedn.-Ochyra,
Sawicki,
Ochyra, Szczecińska & Plášek]
9 R. hespericum Sérgio, J.Muñoz & Ochyra [Codriophorus hespericus (Sérgio, J.Muñoz
& Ochyra) Bedn.-Ochyra & Ochyra]
subgenus Ellipticodryptodon (Vilh.) Bedn.-Ochyra
& Ochyra [Bucklandiella Roiv.]
10 R. affine (F.Weber & D.Mohr) Lindb. [Bucklandiella affinis (Schleich. ex F.Weber
& D.Mohr) Bedn.-Ochyra & Ochyra]
11 R. ellipticum (Turner) Bruch & Schimp. [Bucklandiella elliptica (Turner) Bedn.-Ochyra
& Ochyra]
12 R. heterostichum (Hedw.) Brid. [Bucklandiella heterosticha (Hedw.) Bedn.Ochyra & Ochyra]
13 R. himalayanum (Mitt.) A.Jaeger [Bucklandiella himalayana (Mitt.) Bedn.Ochyra & Ochyra]
14 R. lamprocarpum (Müll.Hal.) A.Jaeger
[Bucklandiella lamprocarpa (Müll.Hal.)
Bedn.-Ochyra & Ochyra]
15 R. lusitanicum Ochyra & Sérgio [Bucklandiella
lusitanica (Ochyra & Sérgio) Bedn.Ochyra & Ochyra]
16 R. macounii Kindb. [Bucklandiella macounii
(Kindb.) Bedn.-Ochyra & Ochyra]
a subsp. alpinum (E.Lawton) Frisvoll [Bucklandiella macounii subsp. alpina
(E.Lawton) Bedn.-Ochyra & Ochyra]
b subsp. macounii
17 R. microcarpon (Hedw.) Brid. [Bucklandiella
microcarpa (Hedw.) Bedn.-Ochyra &
Ochyra]
18 R. nivale (Köckinger, Bedn.-Ochyra & Ochyra)
Köckinger [Bucklandiella nivalis Köckinger, Bedn.-Ochyra & Ochyra]317
19 R. obtusum (Brid.) Brid. [Bucklandiella
obtusa (Brid.) Bedn.-Ochyra & Ochyra]
20 R. sudeticum (Funck) Bruch & Schimp. [Bucklandiella sudetica (Funck) Bedn.-Ochyra
& Ochyra]
Grimmioideae Broth.
120 Coscinodon Spreng.
1 C. cribrosus (Hedw.) Spruce [Grimmia cribrosa
Hedw.]
2 C. horridus (Muñoz & H.Hespanhol) Hugonnot, R.D.Porley & Ignatov [Grimmia
horrida Muñoz & H.Hespanhol]318
The treatment of Racomitrium follows Larraín et al. (2013), who used molecular evidence for a broad treatment of the genus. Sawicki et al. (2015) used
mitochondrial DNA to support narrower genera. However, they used only ten species of Racomitrium s.lat. (two from each segregate genus), and the results
did not contradict the previous treatment. Indeed, significant problems remain (e.g. Bucklandiella sensu Ochyra is not monophyletic) and until Racomitrium
s.lat. is more completely treated, we prefer to retain Racomitrium in a broad sense.
317
Described as Bucklandiella nivalis by Köckinger, Bednarek-Ochyra, et al. (2007) but listed as Racomitrium nivale by Köckinger, Suanjak, et al. (2008).
318
Grimmia horrida was described by Muñoz et al. (2009), and transferred to Coscinodon by Hugonnot et al. (2018).
JOURNAL OF BRYOLOGY
3
4
121
319
C. humilis Milde319
C. monchiquensis R.D.Porley, Ochyra &
Ignatova320
Grimmia Hedw. [Dryptodon Brid., Guembelia
Hampe, Hydrogrimmia (I.Hagen) Loeske,
Orthogrimmia (Schimp.) Ochyra & Żarnowiec, Streptocolea (I.Hagen) Ochyra &
Żarnowiec]321
1 G. alpestris (F.Weber & D.Mohr) Schleich.
[Orthogrimmia alpestris (F.Weber &
D.Mohr) Ochyra & Żarnowiec]
2 G. anodon Bruch & Schimp.
3 G. anomala Hampe ex Schimp.
4 G. arenaria Hampe [Orthogrimmia arenaria
(Hampe) Ochyra & Żarnowiec]322
5 G. atrata Miel. ex Hornsch. [Streptocolea
atrata (Miel. ex Hornsch.) Ochyra &
Żarnowiec]
6 G. caespiticia (Brid.) Jur. [Orthogrimmia caespiticia (Brid.) Ochyra & Żarnowiec]
7 G. capillata De Not.323
8 G. crinita Brid.
9 G. crinitoleucophaea Cardot [Grimmia poecilostoma Cardot & Sebille]324
10 G. curviseta Bouman325
11 G. decipiens (Schultz) Lindb.
12 G. dissimulata E.Maier
13 G. donniana Sm. [Orthogrimmia donniana
(Sm.) Ochyra & Żarnowiec]
14 G. elatior Bruch ex Bals.-Criv. & De Not.
15 G. elongata Kaulf.
16 G. funalis (Schwägr.) Bruch & Schimp.
17 G. fuscolutea Hook.
18 G. hartmanii Schimp.
19 G. incurva Schwägr.
20 G. laevigata (Brid.) Brid. [Guembelia laevigata (Brid.) Ochyra & Żarnowiec]
21 G. lisae De Not.
45
22 G. longirostris Hook. [Guembelia longirostris
(Hook.) Ochyra & Żarnowiec]
23 G. meridionalis (Müll.Hall.) E.Maier [Grimmia
trichophylla
var.
meridionalis
326
Müll.Hall.]
24 G. mollis Bruch & Schimp. [Hydrogrimmia
mollis (Bruch & Schimp.) Loeske]
25 G. montana Bruch & Schimp. [Orthogrimmia
montana (Bruch & Schimp.) Ochyra &
Żarnowiec]
26 G. muehlenbeckii Schimp.
27 G. nutans Bruch [Grimmia meteorae
C.C.Towns.]
28 G. orbicularis Bruch ex Wilson [Dryptodon
orbicularis (Bruch ex Wilson) Ochyra
& Żarnowiec]
29 G. ovalis (Hedw.) Lindb. [Guembelia ovalis
(Hedw.) Müll.Hal.]
30 G. plagiopodia Hedw.
31 G. pulvinata (Hedw.) Sm.
32 G. ramondii (Lam. & DC.) Margad. [Dryptodon patens (Hedw.) Brid., Grimmia
curvata (Brid.) De Sloover]
33 G. reflexidens Müll.Hal. [Grimmia sessitana
De Not., Orthogrimmia sessitana (De
Not.) Ochyra & Żarnowiec]327
34 G. teretinervis Limpr.
35 G. tergestina Tomm. ex Bruch & Schimp.
[Grimmia tergestina var. tergestinoides
(Culm.) Podp., Guembelia tergestina
(Tomm. ex Bruch & Schimp.) Buyss.]328
36 G. torquata Drumm.
37 G. trichophylla Grev. [Grimmia britannica
A.J.E.Sm.]
38 G. triformis Carestia & De Not. [Orthogrimmia triformis (Carestia & De Not.)
Ochyra & Żarnowiec]329
39 G. ungeri Jur.330
40 G. unicolor Hook.
A molecular study by Ignatova, Kuznetsova, et al. (2008) showed that Coscinodon humilis is a European endemic known only from the Alps, whereas plants
with this name in east Asia belong to Coscinodon yukonensis Hastings.
Coscinodon monchiquensis was described from the Algarve, Portugal, by Ignatov, Porley, et al. (2018).
321
Although Ochyra, Żarnowiec, et al. (2003) presented a morphological scheme comprising seven distinct genera, based largely on the traditional subgenera
of Grimmia, we choose to treat it in the broad sense. There is molecular support for the paraphyletic nature of Grimmia (Streiff 2006; Hernández-Maqueda
et al. 2008), but there remain some significant incongruities between plastid and nuclear ITS and the morphological schemes. The evolutionary history of
Grimmia is complex and further sampling and better phylogenetic markers are needed.
322
Grimmia arenaria was synonymised with Grimmia donniana by Maier (2010) but treated as a distinct species in Muñoz et al. (2015). Molecular investigations indicate that it should be accepted as a species (D. Callaghan, pers. comm. 2019).
323
Grimmia capillata was synonymised with G. crinita by Maier (2010) but this view has not gained acceptance, since molecular studies support it at specific
level. It is recognised at species level in Muñoz et al. (2015).
324
Grimmia poecilostoma was synonymised with Grimmia crinitoleucophaea by Muñoz et al. (2015). However, the synonymy of G. crinitoleucophaea with
Grimmia tergestina (Maier 2010) has not been widely accepted, on morphological grounds.
325
Grimmia curviseta was synonymised with G. orbicularis by Maier (2010) but this is disputed; Rodríguez-Romero et al. (2016) used DNA sequencing to conclude that it is an allopolyploid derivative with G. orbicularis as one of its progenitors.
326
Grimmia meridionalis was elevated to the rank of species by Maier (2002), but, unlike Grimmia dissimulata, not accepted by Hill et al. (2006). However, its
status as a species has subsequently been supported by molecular evidence (Streiff 2006).
327
Taxonomic, nomenclatural and misidentification issues surround Grimmia reflexidens. The situation in Europe remains unresolved but we retain Grimmia
reflexidens for the present.
328
Grimmia tergestina var. tergestinoides seems distinct in central Europe. More work is needed to determine its taxonomic value.
329
Grimmia triformis was synonymised with Grimmia donniana by Maier (2010) but differs in several respects, and was accepted as a species by Muñoz et al.
(2015).
330
Grimmia ungeri was listed as a synonym of Grimmia montana by Maier (2010). Muñoz and Pando (2000) listed Grimmia ungeri from several countries,
including Spain and Italy, but it was not included by Muñoz et al. (2015). Specimens from Mexico were redetermined as Grimmia montana
320
46
122
N. G. HODGETTS ET AL.
Schistidium Bruch & Schimp.
1 S. abrupticostatum (Bryhn) Ignatova &
H.H.Blom [Schistidium platyphyllum var.
abrupticostatum (Bryhn) H.H.Blom]331
2 S. agassizii Sull. & Lesq.
3 S. andreaeopsis (Müll.Hal.) Laz.332
4 S. apocarpum (Hedw.) Bruch & Schimp.
5 S. atrichum
(Müll.Hal.
&
Kindb.)
333
W.A.Weber
6 S. atrofuscum (Schimp.) Limpr.
7 S. boreale Poelt
8 S. brunnescens Limpr.
a subsp. brunnescens
b subsp. griseum (Nees & Hornsch.) H.H.Blom
9 S. bryhnii I.Hagen
10 S. canadense (Dupr.) Ignatova & H.H.Blom334
11 S. confertum (Funck) Bruch & Schimp.
12 S. confusum H.H.Blom
13 S. convergens J.Guerra & M.J.Cano335
14 S. crassipilum H.H.Blom
15 S. crenatum H.H.Blom
16 S. dupretii (Thér.) W.A.Weber
17 S. echinatum Ignatova & H.H.Blom336
18 S. elegantulum H.H.Blom
a subsp. elegantulum
b subsp. wilsonii H.H.Blom
19 S. flaccidum (De Not.) Ochyra
20 S. flexipile (Lindb. ex Broth.) G.Roth
21 S. frigidum H.H.Blom
a var. frigidum
b var. havaasii H.H.Blom
22 S. frisvollianum H.H.Blom
23 S. grande Poelt
24 S. grandirete H.H.Blom
25 S. helveticum (Schkuhr) Deguchi [Schistidium
singarense (Schiffn.) Laz.]
26 S. holmenianum Steere & Brassard
27 S. lancifolium (Kindb.) H.H.Blom
28 S. marginale H.H.Blom, Bedn.-Ochyra &
Ochyra337
29 S. maritimum (Sm. ex R.Scott) Bruch &
Schimp.
a subsp. maritimum
b subsp. piliferum (I.Hagen) B.Bremer
30 S. obscurum H.H.Blom, Köckinger &
Ignatova338
31 S. occidentale (E.Lawton) S.P.Churchill
32 S. papillosum Culm.
33 S. platyphyllum (Mitt.) H.Perss.
34 S. poeltii H.H.Blom
35 S. pruinosum (Wilson ex Schimp.) G.Roth
36 S. pulchrum H.H.Blom
37 S. recurvum H.H.Blom
38 S. rivulare (Brid.) Podp.
39 S. robustum (Nees & Hornsch.) H.H.Blom
40 S. scandicum H.H.Blom
41 S. sibiricum Ignatova & H.H.Blom339
42 S. sinensiapocarpum (Müll.Hal.) Ochyra340
43 S. sordidum I.Hagen
44 S. spinosum H.H.Blom & Lüth
45 S. strictum (Turner) Loeske ex Mårtensson
46 S. subconfertum (Broth.) Deguchi341
47 S. subflaccidum (Kindb.) H.H.Blom342
48 S. subjulaceum H.H.Blom
49 S. submuticum H.H.Blom
a subsp. arcticum H.H.Blom
b subsp. submuticum
50 S. succulentum Ignatova & H.H.Blom343
51 S. tenerum (J.E.Zetterst.) Nyholm
52 S. tenuinerve Ignatova & H.H.Blom344
53 S. trichodon (Brid.) Poelt
a var. nutans H.H.Blom
b var. trichodon
54 S. umbrosum (J.E.Zetterst.) H.H.Blom
55 S. venetum H.H.Blom
Hedwigiales Ochyra
Hedwigiaceae Schimp.
123
Braunia Bruch & Schimp. [Hedwigidium Bruch &
Schimp.]
(Delgadillo-Moya 2015). On the other hand, it differs from Grimmia montana in some respects (J. Muñoz, pers. comm. 2017). Although we retain it in the
checklist for now, Grimmia ungeri is taxonomically dubious and may ultimately prove to be synonymous with Grimmia montana.
Schistidium abrupticostatum is not very close to Schistidium platyphyllum according to molecular studies by Ignatova, Blom, et al. (2010).
332
Schistidium andreaeopsis is an Asian Arctic species recorded more recently from the European Arctic (Blom et al. 2006).
333
Schistidium atrichum is principally a North American species with occurrences in France reported by Chavoutier and Hugonnot (2013).
334
Schistidium canadense is a North American species with a single European record from Russia (Karelia) (Ignatova, Blom, et al. 2010).
335
Schistidium convergens was recently described as a new species from southern Spain and Morocco (Guerra, Jiménez-Martínez, Cano, Alonso, et al. 2019).
336
Schistidium echinatum was described from Russia by Ignatova, Blom, et al. (2010), and has also now been found in France and Austria.
337
Schistidium marginale was described from Austria by Blom et al. (2016), and has since been found in several montane regions in Europe.
338
Schistidium obscurum was described from Austria by Ignatova, Blom, et al. (2010). Elsewhere in Europe it has also been found in Switzerland, the Caucasus
and Svalbard.
339
Schistidium sibiricum was described from Russia by Ignatova, Blom, et al. (2010).
340
Schistidium sinensiapocarpum was first reported in Europe from the Caucasus (Blom et al. 2006), and also occurs in the Austrian Alps (Köckinger, Suanjak,
et al. 2008).
341
Schistidium subconfertum is primarily an Asian species. However, a herbarium specimen was recently discovered from the Italian Alps (Italy: Stilfserjochhöhe, 26 August 1909, Glowacki, GJO), and examined by H. Blom. This species is here published new to Europe.
342
Schistidium subflaccidum was reported from the Caucasus in Russia (Blom et al. 2006), and has subsequently been found in mountainous regions of other
parts of Europe.
343
Schistidium succulentum was described from Russia by Ignatova, Blom, et al. (2010).
344
Schistidium tenuinerve was described from Asiatic Russia (Ignatova, Blom, et al. 2010), but was recently also recorded from European Russia (E. Ignatova
pers. comm. 2018).
331
JOURNAL OF BRYOLOGY
1
2
124
B. alopecura (Brid.) Limpr.
B. imberbis (Sm.) N.Dalton & D.G.Long [Hedwigidium integrifolium auct., non
(P.Beauv.) Dixon, Hedwigia integrifolia
auct., non P.Beauv.]345
Hedwigia P.Beauv.
1 H. ciliata (Hedw.) P.Beauv.
2 H. emodica Hampe ex Müll.Hal. [Hedwigia ciliata
var. leucophaea Bruch & Schimp.]346
3 H. mollis Ignatova, Ignatov & Fedosov347
4 H. nemoralis Ignatova, Ignatov & Fedosov348
5 H. stellata Hedenäs
6 H. striata (Bruch & Schimp.) John Whitehead
& Fergusson ex Hobk. & Porritt349
Bartramiales D.Quandt, N.E.Bell & M.Stech
Bartramiaceae Schwägr.
Conostomoideae D.G.Griffin & W.R.Buck
125 Conostomum Sw.
1 C. tetragonum (Hedw.) Lindb.
Bartramioideae D.G.Griffin & W.R.Buck
126 Anacolia Schimp.
1 A. menziesii (Turner) Paris
2 A. webbii (Mont.) Schimp.
127
345
Bartramia Hedw.
section Bartramia
1 B. halleriana Hedw.
2 B. pomiformis Hedw.
section Pyridium Müll.Hal.
3 B. breviseta Lindb.
4 B. ithyphylla Brid.
5 B. subulata Bruch & Schimp.
section Strictidium Müll.Hal.
6 B. aprica Müll.Hal. [Bartramia rosamrosiae
Damayanti, J.Muñoz, J.-P.Frahm &
D.Quandt, Bartramia stricta auct.
eur., non Brid.]350
7 B. laevisphaera (Taylor) Müll.Hal. [Anacolia
laevisphaera (Taylor) Flowers]351
128
47
Plagiopus Brid.
1 P. oederianus (Sw.) H.A.Crum & L.E.Anderson
a var. alpinus (Schwägr.) Ochyra352
b var. oederianus
Breutelioideae D.G.Griffin & W.R.Buck
129 Breutelia (Bruch & Schimp.) Schimp.
1 B. azorica (Mitt.) Cardot
2 B. chrysocoma (Hedw.) Lindb.
130
Philonotis Brid.
section Bartramidula (Bruch & Schimp.) Mitt.
1 P. cernua (Wilson) D.G.Griffin & W.R.Buck
section Philonotula (Schimp.) A.Jaeger
2 P. rigida Brid.
section Homomorphae (Kindb.) Ochyra
3 P. capillaris Lindb. [Philonotis arnellii
Husn.]353
4 P. falcata (Hook.) Mitt.354
5 P. hastata (Duby) Wijk & Margad.
6 P. marchica (Hedw.) Brid.
7 P. uncinata (Schwägr.) Brid.
section Philonotis
8 P. caespitosa Jur.
9 P. calcarea (Bruch & Schimp.) Schimp.
10 P. fontana (Hedw.) Brid.
11 P. seriata Mitt.
12 P. tomentella Molendo
13 P. yezoana Besch. & Cardot355
Splachnales Ochyra
Splachnaceae Grev. & Arn.
Taylorioideae Broth.
131 Tayloria Hook.
1 T. acuminata Hornsch.
2 T. froelichiana (Hedw.) Mitt. ex Broth.
3 T. hornschuchii (Grev. & Arn.) Broth.
4 T. lingulata (Dicks.) Lindb.
5 T. rudolphiana (Garov.) Bruch & Schimp.
6 T. serrata (Hedw.) Bruch & Schimp.
7 T. splachnoides (Schleich. ex Schwägr.) Hook.
8 T. tenuis (Dicks.) Schimp.
Dalton et al. (2012) found that European specimens identified as Hedwigidium integrifolium differed from the type material of Hedwigia integrifolia
P.Beauv., and were considered to belong to Braunia imberbis. De Luna (2016) reported that Braunia imberbis, along with some other (non-European)
species, group together in a different clade, Hedwigidium. Until further research is done on this group, the species is here retained in Braunia.
346
Hedenäs (1994) suggested that Hedwigia emodica was a synonym of Hedwigia ciliata var. leucophaea. Buchbender et al. (2014) later proved the hybrid
origin of some accessions referable to this taxon. Ignatova, Kuznetsova, et al. (2016) recognised the taxon at species level, as Hedwigia emodica, but doubts
persist that all European material is the same as the plants from Russia (or the Himalayan type of Hedwigia emodica).
347
Ignatova, Kuznetsova, et al. (2016) described Hedwigia mollis as part of their study of the genus in Russia.
348
Hedwigia nemoralis was described from Russia by Ignatova, Kuznetsova, et al. (2016).
349
Hedwigia striata was a neglected taxon returned to specific rank by Buchbender et al. (2014).
350
Damayanti et al. (2012) described Bartramia rosamrosiae for the European plants after finding that Bartramia stricta was restricted to South America. This
was synonymised with Bartramia aprica by Müller (2014). However, recent (as yet unpublished) work by Neumann, Muñoz and Quandt suggests that Bartramia aprica and Bartramia rosamrosiae are indeed distinct species.
351
Molecular analysis revealed that Bartramia laevisphaera belongs to Bartramia section Strictidium rather than to Anacolia (Damayanti et al. 2012).
352
There is disagreement over the status of Plagiopus oederianus var. alpinus: it may be nothing more than a high-altitude form of the species, but is retained
for the present.
353
Philonotis capillaris is the correct name for Philonotis arnellii (Koponen and Isoviita 2010).
354
Philonotis falcata is an Asian species reported from the Caucasus new to Europe by Ignatov, Fedosov, et al. (2010).
355
Philonotis yezoana is an Asian species reported from Finland new to Europe by Ulvinen and Kypärä (2016).
48
N. G. HODGETTS ET AL.
Splachnoideae Broth.
132 Aplodon R.Br.
1 A. wormskioldii (Hornem.) R.Br.
133
Splachnum Hedw.
1 S. ampullaceum Hedw.
2 S. luteum Hedw.
3 S. melanocaulon (Wahlenb.) Schwägr.
4 S. pensylvanicum (Brid.) Grout ex H.A.Crum
5 S. rubrum Hedw.
6 S. sphaericum Hedw.
7 S. vasculosum Hedw.
134
Tetraplodon Bruch & Schimp.
1 T. angustatus (Hedw.) Bruch & Schimp.
2 T. blyttii Frisvoll
3 T. mnioides (Hedw.) Bruch & Schimp.
4 T. pallidus I.Hagen
5 T. paradoxus (R.Br.) I.Hagen
6 T. urceolatus (Hedw.) Bruch & Schimp.
135
Voitia Hornsch.
1 V. hyperborea Grev. & Arn.
2 V. nivalis Hornsch.
Meesiaceae Schimp.
136 Amblyodon P.Beauv.
1 A. dealbatus (Hedw.) P.Beauv.
137
Leptobryum (Bruch & Schimp.) Wilson
1 L. pyriforme (Hedw.) Wilson
138
Meesia Hedw
1 M. hexasticha (Funck) Bruch
2 M. longiseta Hedw.
3 M. triquetra (L. ex Jolycl.) Ångstr.
4 M. uliginosa Hedw.
139
Paludella Brid.
1 P. squarrosa (Hedw.) Brid.
Bryales Limpr.
Bryaceae Schwägr.356
356
140
Anomobryum Schimp.
1 A. apiculatum (Schwägr.) D.Bell & Holyoak
[Bryum apiculatum Schwägr., Gemmabryum apiculatum (Schwägr.)
J.R.Spence & H.P.Ramsay, Osculatia
apiculata (Schwägr.) Ochyra, Plášek
& Bedn.-Ochyra]357
2 A. concinnatum (Spruce) Lindb.358
3 A. julaceum (Schrad. ex P.Gaertn., E.Mey &
Scherb.) Schimp.
4 A. lusitanicum (I.Hagen) Thér.
5 A. notarisii (Mitt.) D.Bell. & Holyoak [Brachymenium notarisii (Mitt.) A.J.Shaw]359
141
Brachymenium Schwägr.
1 B. paradoxum (Herzog) A.J.Shaw
142
Bryum Hedw. [Gemmabryum J.R.Spence &
H.R.Ramsay, Osculatia De Not.]360
1 B. argenteum Hedw.*
2 B. bavaricum Warnst. [Anomobryum bavaricum
(Warnst.)
Holyoak
&
Köckinger]*361
3 B. blindii Bruch & Schimp.
4 B. canariense Brid. [Rosulabryum canariense
(Brid.) Ochyra]362
5 B. caucasicum (Schimp. ex Broth.) C.J.Cox &
Hedd.*
6 B. demaretianum Arts [Gemmabryum demaretianum (Arts) J.R.Spence, Osculatia
demaretiana (Arts) Ochyra, Plášek &
Bedn.-Ochyra]
7 B. dichotomum Hedw. [Bryum barnesii
J.B.Wood ex Schimp., Gemmabryum
barnesii (J.B.Wood ex Schimp.)
J.R.Spence,
Osculatia
barnesii
(J.B.Wood ex Schimp.) Ochyra,
Plášek & Bedn.-Ochyra, Osculatia
bicolor (Dicks.) Ochyra, Plášek &
Bedn.-Ochyra, Gemmabryum dichotomum
(Hedw.)
J.R.Spence
&
H.P.Ramsay, Osculatia dichotoma
See note on Bryaceae in introduction.
The only confirmed European record of Bryum apiculatum appears to be a specimen from Tenerife (leg. E. Bourgeon s.n., BM), which was reidentified by
Ochi (1972). That specimen may nevertheless need critical reappraisal since Ochi did not record Bryum gemmiparum which is locally frequent on Tenerife,
and there have been no subsequent reports of Bryum apiculatum occurring there. Holyoak (2009) and in Blockeel et al. (2014) identified plants from Cornwall as this species and two additional localities have since been found in the same county. However, data from DNA barcoding markers and preliminary
analyses of a substantial genomic dataset place Cornish specimens (Holyoak 09-42 and 07-15) in Bryum s.str., close to Bryum dichotomum, differing markedly from a specimen of Bryum apiculatum from Puerto Rico (R.D. Worthington 35254). The Cornish plants thus represent an undescribed taxon allied to
Bryum dichotomum, an account of which is in preparation.
358
Holyoak and Köckinger (2010) supported treatment of Anomobryum concinnatum as a distinct species from Anomobryum julaceum based on morphological characters. Data from DNA barcoding markers (D. Bell et al., in litt.) has subsequently shown that Anomobryum concinnatum and Anomobryum julaceum
form separate clades.
359
Recent molecular work has shown that Brachymenium notarisii is closely related to Anomobryum lusitanicum, and is therefore probably best treated in that
genus (D.Bell et al. in litt.).
360
Species with adequate molecular data confirming their position in Bryum s.str. are annotated with an asterisk (*). The remaining species lack sufficiently
informative data, but there are now relatively few remaining ‘orphan’ species in Bryum, several more having been transferred to Ptychostomum or Imbribryum on the basis of very recent DNA evidence.
361
Holyoak and Köckinger (2010) placed Anomobryum bavaricum in Anomobryum on the basis of morphology, but molecular data places it in Bryum s.str.
(D.Bell et al., in litt.).
362
Results from DNA barcoding and nuclear genomic datasets (D. Bell et al., in litt. ) show that Bryum canariense does not belong in Bryum s.str., but is part of a
near-basal clade of Bryaceae allied to Brachymenium s.str. and Leptostomopsis. However, it is retained in Bryum here pending further investigation.
357
JOURNAL OF BRYOLOGY
8
9
10
11
12
13
14
15
16
17
18
19
20
363
(Hedw.) Ochyra, Plášek & Bedn.Ochyra]*363
B. dixonii Cardot ex W.E.Nicholson
B. dyffrynense Holyoak*
B. gemmiferum R.Wilczek & Demaret [Gemmabryum gemmiferum (R.Wilczek &
Demaret) J.R.Spence, Osculatia gemmifera (R.Wilczek & Demaret)
Ochyra, Plášek & Bedn.-Ochyra]*
B. gemmilucens R.Wilczek & Demaret [Gemmabryum gemmilucens (R.Wilczek &
Demaret) J.R.Spence, Osculatia gemmilucens (R.Wilczek & Demaret)
Ochyra, Plášek & Bedn.-Ochyra]*
B. gemmiparum De Not. [Imbribryum gemmiparum (De Not.) J.R.Spence]*
B. kikuyense (Broth. & Thér.) N.Pedersen
[Brachymenium philonotula Broth.,
Bryum philonotulum Hampe nom.
inval., non Bryum philonotulum
Müll.Hal.]*364
B. klinggraeffii Schimp. [Gemmabryum klinggraeffii (Schimp.) J.R.Spence &
H.P.Ramsay, Osculatia klinggraeffii
(Schimp.) Ochyra, Plášek & Bedn.Ochyra ]365
B. marratii Hook.f. & Wilson
B. oblongum Lindb.366
B. radiculosum Brid. [Gemmabryum radiculosum (Brid.) J.R.Spence & H.P.
Ramsay, Osculatia radiculosa (Brid.)
Ochyra, Plášek & Bedn.-Ochyra]*
B. reyeri Breidl.367
B. riparium I.Hagen
B. ruderale Crundw. & Nyholm [Gemmabryum ruderale (Crundw. & Nyholm)
J.R.Spence,
Osculatia
ruderalis
21
22
23
24
143
49
(Crundw. & Nyholm) Ochyra, Plášek
& Bedn.-Ochyra]*
B. sauteri Bruch & Schimp. [Osculatia sauteri
(Bruch & Schimp.) Ochyra, Plášek &
Bedn.-Ochyra]
B. valparaisense Thér. [Gemmabryum valparaisense (Thér.) J.R.Spence, Osculatia valparaisense (Thér.) Ochyra,
Plášek & Bedn.-Ochyra]
B. versicolor A.Braun ex Bruch & Schimp.368
B. violaceum Crundw. & Nyholm [Gemmabryum violaceum (Crundw. & Nyholm)
J.R.Spence,
Osculatia
violacea
(Crundw. & Nyholm) Ochyra, Plášek
& Bedn.-Ochyra]*
Imbribryum Pedersen369
1 I. alpinum (Huds. ex With.) N.Pedersen
[Bryum alpinum Huds. ex With.]
2 I. mildeanum (Jur.) J.R.Spence [Bryum mildeanum Jur.]
3 I. miniatum (Lesq.) J.R.Spence [Bryum miniatum Lesq.]370
4 I. muehlenbeckii (Bruch & Schimp.) N.Pedersen [Bryum muehlenbeckii Bruch &
Schimp.]371
5 I. subapiculatum (Hampe) D.Bell & Holyoak
[Bryum subapiculatum Hampe, Gemmabryum subapiculatum (Hampe)
J.R.Spence & H.P.Ramsay, Osculatia
subapiculata (Hampe) Ochyra, Plášek
& Bedn.-Ochyra]372
6 I. tenuisetum (Limpr.) D.Bell & Holyoak
[Bryum tenuisetum Limpr., Gemmabryum tenuisetum (Limpr.) J.R.Spence
& H.P. Ramsay, Osculatia tenuiseta
(Limpr.) Ochyra, Plášek & Bedn.Ochyra]373
Bryum barnesii is usually regarded as a distinct species in Central Europe, but many plants and populations intermediate between Bryum barnesii and
Bryum dichotomum occur (Holyoak 2003).
Molecular data show that Brachymenium philonotula Broth. should be placed in Bryum rather than Brachymenium, but since the name Bryum philonotula
Müll.Hal. (≡ Pohlia philonotula (Müll.Hal.) Broth.) had already been used, the valid name for the species becomes Bryum kikuyense (Pedersen and Hedenäs
2005).
365
According to molecular evidence from Holyoak and Pedersen (2007), Bryum klinggraeffii could possibly be placed in Imbribryum or its own genus, but is
retained in Bryum for the present.
366
Preliminary molecular data suggest that Bryum oblongum may belong to Ptychostomum s. lat, but further study is needed to confirm this.
367
Bryum reyeri is only known from the eastern Italian Alps (type), Austria and Germany (Limpricht 1895; Grims 1999; Meinunger and Schröder 2007; Köckinger, Suanjak, et al. 2008). Provisional data from a nuclear DNA dataset (D. Bell et al., in litt.) imply it is part of a clade basal to the radiation of Ptychostomum, as circumscribed here.
368
Bryum versicolor was included as a synonym of Bryum dichotomum by Hill et al. (2006) but this is a widely recognised taxon, so it is reinstated pending
further studies.
369
Imbribryum was recognised at generic level by Holyoak and Pedersen (2007), and new molecular data from a substantial nuclear genomic dataset supports
this view (D. Bell et al., in litt.).
370
Imbribryum miniatum is placed in Imbribryum based on molecular data from specimens from both the Faeroes and the U.S.A.
371
Holyoak (in Hill et al. 2006) synonymised Bryum muehlenbeckii with Bryum alpinum because it appeared to be only a form resulting from growth underwater or beneath snow, and there are numerous intermediate specimens. However, DNA sequence data for Imbribryum muehlenbeckii from three exemplars from different localities define a single clade that is well separated from Imbribryum alpinum. Hence, they are regarded here as separate species in
which occasional apparent morphological intergradation is due to the occurrence of phenotypes (‘expressions’) that match the appearance of the other
taxon. Additional molecular data suggest that it may be better treated in Ptychostomum, but further work is needed before this can be confirmed.
372
DNA barcode data show Bryum subapiculatum strongly supported in a clade with Imbribryum alpinum (D. Bell et al., in litt.), and it is therefore transferred to
Imbribryum.
373
Along with Bryum subapiculatum, DNA barcoding places Bryum tenuisetum in a clade with Imbribryum alpinum (D. Bell et al., in litt.), so it is here transferred
to Imbribryum.
364
50
144
374
N. G. HODGETTS ET AL.
Ptychostomum
Hornsch.
[Plagiobryoides
J.R.Spence, Plagiobryum Lindb., Rosulabryum J.R.Spence]374
1 P. arcticum (R.Br.) J.R.Spence ex Holyoak &
N.Pedersen [Bryum arcticum (R.Br.)
Bruch & Schimp.]
2 P. austriacum (Köckinger, Holyoak &
Suanjak) D.Bell & Holyoak [Bryum
austriacum Köckinger, Holyoak &
Suanjak]375
3 P. bornholmense (Wink. & R.Ruthe) Holyoak
& N.Pedersen [Bryum bornholmense
Wink. & R.Ruthe, Osculatia bornholmensis (Wink. & R.Ruthe) Ochyra,
Plášek & Bedn.-Ochyra, Rosulabryum
bornholmense (Wink. & Ruthe)
J.R.Spence]
4 P. calophyllum (R.Br.) J.R.Spence [Bryum
calophyllum R.Br.]376
5 P. capillare (Hedw.) Holyoak & N.Pedersen
[Rosulabryum
capillare
(Hedw.)
J.R.Spence, Bryum capillare Hedw.]
6 P. cellulare (Hook.) D.Bell & Holyoak
[Bryum cellulare Hook., Plagiobryoides
cellularis (Hook.) J.R.Spence]
7 P. cernuum (Hedw.) Hornsch. [Bryum uliginosum (Brid.) Bruch & Schimp.]
8 P. compactum Hornsch. [Bryum algovicum
Sendtn. ex Müll.Hal., Bryum compactum (Hornsch.) Kindb.]
a var. compactum
b var. rutheanum (Warnst.) Holyoak &
N.Pedersen [Bryum algovicum
var.
rutheanum
(Warnst.)
Crundw.]
9
10
11
12
13
14
15
16
17
18
19
20
P. creberrimum (Taylor) J.R.Spence &
H.P.Ramsay [Bryum creberrimum
Taylor]
P. cryophilum (Mårtensson) J.R.Spence
[Bryum cryophilum Mårtensson]377
P. cyclophyllum (Schwägr.) J.R.Spence [Bryum
cyclophyllum (Schwägr.) Bruch &
Schimp.]
P. demissum (Hook.) Holyoak & N.Pedersen
[Plagiobryum
demissum
(Hook.)
Lindb.]
P. donianum (Grev.) Holyoak & N.Pedersen
[Bryum donianum Grev., Rosulabryum
donianum (Grev.) Ochyra]
P. elegans (Nees) D.Bell & Holyoak [Bryum
elegans Nees, Rosulabryum elegans
(Nees) Ochyra]378
P. funkii (Schwägr.) J.R.Spence [Bryum funkii
Schwägr.]379
P. imbricatulum (Müll.Hal.) Holyoak & N.Pedersen [Bryum caespiticium Hedw., Gemmabryum
caespiticium
(Hedw.)
J.R.Spence,
Osculatia
caespiticia
(Hedw.) Ochyra, Plášek & Bedn.-Ochyra]
P. inclinatum (Sw. ex Brid.) J.R.Spence
[Bryum archangelicum Bruch &
Schimp., Ptychostomum archangelicum (Bruch & Schimp.) J.R.Spence]380
P. intermedium (Brid.) J.R.Spence [Bryum
intermedium (Brid.) Blandow]381
P. knowltonii (Barnes) J.R.Spence [Bryum
knowltonii Barnes]382
P. kunzei (Hornsch.) J.R. Spence [Bryum
kunzei
Hornsch.,
Gemmabryum
kunzei
(Hornsch.)
J.R.Spence,
Ptychostomum was recognised at generic level by Holyoak and Pedersen (2007). There are no molecular data on the type species of Rosulabryum, which is
Rosulabryum albolimbatum (Hampe) J.R.Spence from Australia. Recent analyses of a nuclear genomic dataset including the rather similar Rosulabryum
billardierei (Schwägr.) J.R.Spence from the Southern Hemisphere (type from Tasmania), based on a specimen from Ecuador, imply that it forms part of
Brachymenium s.str. Hence, further study is desirable before placing European and N. American species in Rosulabryum.
375
Bryum austriacum was described from the Alps by Köckinger, Holyoak, et al. (2013). DNA barcode data places it in Ptychostomum (D. Bell et al., in litt.).
376
Results from DNA barcoding (D. Bell et al. in litt.) show that Bryum calophyllum belongs in Ptychostomum.
377
DNA barcoding results place Bryum cryophilum in Ptychostomum (D. Bell et al., in litt.).
378
Morphological similarities have led authors of numerous floras (e.g. Schimper 1876; Smith 2004; Spence 2014) to place Bryum elegans close alongside
Bryum capillare. Caution induced by lack of molecular data on elegans nevertheless prevented Holyoak and Pedersen (2007) transferring the species to
Ptychostomum alongside Ptychostomum capillare, and its treatment in Bryum was followed by Ros, Mazimpaka, et al. (2013) and Hodgetts (2015).
However, the molecular-phylogenetic study by Guerra, Jiménez-Martínez, Cano, Jiménez-Fernández, (2011) revealed Bryum elegans as closer to Bryum
pallescens (i.e. Ptychostomum pallescens) than other Bryaceae they analysed, justifying the new combination that is made here. Newly generated
nuclear genomic data also place elegans as part of Ptychostomum s.lat., but outside the (sub-)clade containing P. capillare. Spence (2014) has treated
both species within the genus Rosulabryum, but Pedersen et al. (2003) and Shaw (2014) noted that species previously attributed by Spence to Rosulabryum
were placed in two different clades in a phylogenetic tree based on molecular data. For this reason and because of continuing doubts about the affinities of
the type species Rosulabryum albolimbatum, Rosulabryum is treated here as a synonym of the more widely inclusive genus Ptychostomum.
379
Results from DNA barcoding (D. Bell et al., in litt.) show that Bryum funkii belongs in Ptychostomum. The spelling ‘funkii’ was used in Index Muscorum, but it
was changed to ‘funckii’ because it commemorates Heinrich Christian Funck (1771-1839). Although at first sight this appears to be correction of an orthographical error (under ICBN Art. 60.1) and congruent with the orthography of Marsupella funckii and Funckia named in honour of the same botanist, Funk or
Funck himself used both spellings of his name (biographer E. Hertel, pers. comm. to H. Köckinger), so there is no need to change established usage.
380
Ochyra, Lewis Smith, et al. (2008) pointed out that ”the epithet inclinatum is still available for Ptychostomum imbricatum when it is considered a member of
Ptychostomum”. Since Pohlia inclinata Sw. ex Brid. 1803 is the oldest legitimate name available for the species it must therefore be adopted when the
species is transferred to Ptychostomum, whereas its usage in Bryum was blocked by the later homonym Bryum inclinatum (Hedw.) Dicks. of 1801 (now
Distichium inclinatum). Spence (2005) used the epithet inclinatum under Ptychostomum, but this new combination was invalid since the basionym was
incorrectly cited (contrary to ICBN Art. 33.4). Hence, it was not until publication of Spence (2014) that Ptychostomum inclinatum (Sw. ex Brid.)
J.R.Spence was validated in accordance with ICBN Art. 33.4 and Art. 45.1.
381
The placing of Bryum intermedium in Ptychostomum is supported by sequence data from DNA barcoding markers (D. Bell et al., in litt.).
382
The placing of Bryum knowltonii in Ptychostomum is supported by sequence data from DNA barcoding markers (D. Bell et al., in litt.).
JOURNAL OF BRYOLOGY
21
22
23
24
25
26
27
28
383
Osculatia kunzei (Hornsch.) Ochyra,
Plášek & Bedn.-Ochyra]383
P. longisetum (Blandow ex Schwägr.)
J.R.Spence
[Bryum
longisetum
Blandow ex Schwägr.]
P. minii (Podp. ex Guim.) D.Bell & Holyoak
[Bryum minii Podp. ex Guim.]
P. moravicum (Podp.) Ros & Mazimpaka
[Bryum moravicum Podp., Rosulabryum laevifilum (Syed) Ochyra]
P. pallens (Sw. ex anon.) J.R.Spence [Bryum
pallens Sw. ex anon., Bryum sibiricum
Lindb. & Arnell]384
P. pallescens (Schleich. ex Schwägr.) J.R.Spence
[Bryum pallescens Schleich. ex Schwägr.,
Ptychostomum boreale (F.Weber &
D.Mohr) Ochyra & Bedn.-Ochyra]385
P. pseudotriquetrum (Hedw.) J.R.Spence &
H.P.Ramsay ex Holyoak & N.Pedersen
[Bryum neodamense Itzigs., Bryum
pseudotriquetrum (Hedw.) P.Gaertn.
B.Mey. & Scherb. Ptychostomum neodamense (Itzigs.) J.R.Spence]386
a var. bimum (Schreb.) Holyoak & N.Pedersen [Bryum pseudotriquetrum
var. bimum (Schreb.) Lilj.]
b var. pseudotriquetrum
P. rubens (Mitt.) Holyoak & N.Pedersen
[Bryum rubens Mitt., Osculatia rubens
(Mitt.) Ochyra, Plášek & Bedn.-Ochyra,
Rosulabryum rubens (Mitt.) J.R.Spence]
P. salinum (I.Hagen ex Limpr.) J.R.Spence
[Bryum salinum I.Hagen ex Limpr.]387
51
29 P. schleicheri (DC.) J.R.Spence ex D.Bell &
Holyoak [Bryum schleicheri DC.]388
a var. latifolium (Schwägr.) D.Bell &
Holyoak [Bryum schleicheri var.
latifolium (Schwägr.) Schimp.]
b var. schleicheri
30 P. torquescens (Bruch & Schimp.) Ros &
Mazimpaka
[Bryum
torquescens
Bruch & Schimp., Rosulabryum torquescens
(Bruch
&
Schimp.)
J.R.Spence]
31 P. turbinatum (Hedw.) J.R.Spence [Bryum
turbinatum (Hedw.) Turner]389
32 P. warneum (Röhl.) J.R.Spence [Bryum
warneum (Röhl.) Brid.]390
33 P. weigelii (Biehler) J.R.Spence [Bryum weigelii Biehler]391
34 P. wrightii (Sull. & Lesq.) J.R.Spence [Bryum
wrightii Sull. & Lesq.]392
35 P. zieri (Hedw.) Holyoak & N.Pedersen [Plagiobryum zieri (Hedw.) Lindb.]
145
Rhodobryum (Schimp.) Limpr.
1 R. ontariense (Kindb.) Kindb.
2 R. roseum (Hedw.) Limpr.
Mniaceae Schwägr.
Mielichhoferioideae M.Stech & W.Frey
146 Mielichhoferia Nees & Hornsch.
1 M. elongata (Hoppe & Hornsch. ex Hook.)
Hornsch.
2 M. mielichhoferiana (Funck) Loeske
DNA barcoding results suggest that Bryum kunzei should be transferred to Ptychostomum (D. Bell et al., in litt.).
Zolotov (2007a) regarded Bryum sibiricum as possibly ”a good, but very rare species”, with the comment that ”it is rather contrastingly different from
B. pallens which is relatively little variable in gametophytic characters”, all this on the basis of study of a single specimen placed as Bryum sibiricum
from Yakutia because it matched the original description well. Zolotov (2007b) noted it as close Bryum pallens, but ”differs … [in the] very narrow
leaves”. Zolotov made no mention though of the great variability of Ptychostomum pallens revealed in the extensive illustrated monograph of the
species by Podpěra (1973), which included the new combination Bryum pallens subsp. sibiricum (Lindb. & Arnell) Podp. (p. 195), redescribed in detail.
Other forms of Bryum pallens with long narrow leaves and excurrent costa were figured and described in Podpěra’s work, including subsp. prosboreum
Podp. (from arctic Sweden), subsp. acutiusculum Podp. (Alaska) and forma gracile Podp. (Bohemia), as well as other infraspecific taxa intermediate between
these and normal Ptychostomum pallens.
385
Ochyra & Bednarek-Ochyra (2011) demonstrated that Hypnum boreale F.Weber & D.Mohr, 1807, and the subsequent combination Bryum boreale (F.Weber
& D.Mohr) Funck, 1808, are older names for Ptychostomum pallescens than Bryum pallescens Schwägr., 1816. Ochyra and Bednarek-Ochyra (2015) published
a proposal to maintain established usage of the name Bryum pallescens in preference to the unfamiliar and largely forgotten name Bryum boreale and this
desirable initiative is supported here. The combination Ptychostomum boreale (F.Weber & D.Mohr) Ochyra & Bednarek-Ochyra, 2011 was valid, and adopted
as correct by Ros, Mazimpaka, et al. (2013) and Hodgetts (2015). However, pending the response to the proposal by Ochyra and Bednarek-Ochyra (2015),
the established usage should be maintained for Ptychostomum pallescens.
386
Ptychostomum neodamense has been shown by Holyoak and Hedenäs (2006), from molecular and other data, to represent an inconstant phenotype of
Ptychostomum pseudotriquetrum occurring locally in highly calcareous habitats prone to inundation, and connected to it by intermediate forms known
widely in Europe and also in Asia, Alaska and Greenland. Its retention at species rank by Spence (2014) is a consequence of an over-emphasis on morphological data.
387
Bryum salinum is very close to Ptychostomum inclinatum (Holyoak and Pedersen 2007), and this is supported by sequence data from DNA barcoding
markers (D. Bell et al. in litt.).
388
DNA barcoding results and data from a nuclear genomic dataset show that Bryum schleicheri is better placed in Ptychostomum (D. Bell et al., in litt.).
However, relationships between the two varieties remain uncertain, as the two datasets are not directly comparable, and a study is needed to clarify
the position of Ptychostomum schleicheri var. latifolium. As a large, swollen plant with concave leaves, it is the plant that is understood as
Ptychostomum schleicheri in many parts of Europe; var. schleicheri is less well known, sometimes because it is misidentified as Ptychostomum turbinatum.
389
Spence’s placement of Bryum turbinatum in Ptychostomum is supported by molecular evidence (D. Bell et al., in litt.).
390
Spence’s placement of Bryum warneum in Ptychostomum is supported by molecular evidence (D. Bell et al., in litt.).
391
Ptychostomum weigelii has been considered an invalid combination since Spence (2005) cited Sprengel (1807, July) as the author of the basionym name
Bryum weigelii, when the name was in fact published validly by Biehler (1807, May) a couple of months earlier. However, since Spence simply cited the
wrong publication, and did not omit any clear reference to the basionym, this is a correctable error (cf. ICBN Art. 41.8(a); Turland et al. 2018).
392
Bryum wrightii was placed in Ptychostomum by Holyoak and Pedersen (2007), following Pedersen et al. (2003).
384
52
147
148
N. G. HODGETTS ET AL.
Pohlia Hedw.
1 P. andalusica (Höhn.) Broth.
2 P. andrewsii A.J.Shaw
3 P. annotina (Hedw.) Lindb.
4 P. atropurpurea (Wahlenb.) H.Lindb.
5 P. beringiensis A.J.Shaw393
6 P. bolanderi (Lesq.) Broth.
7 P. bulbifera (Warnst.) Warnst.
8 P. camptotrachela (Renauld & Cardot) Broth.
9 P. cruda (Hedw.) Lindb.
10 P. crudoides (Sull. & Lesq.) Broth.
11 P. drummondii (Müll.Hal.) A.L.Andrews
12 P. elongata Hedw.
a var. acuminata (Hornsch.) Huebener
b var. elongata
c var. greenii (Brid.) A.J.Shaw
13 P. erecta Lindb.
14 P. filum (Schimp.) Mårtensson
15 P. flexuosa Harv.
a var. flexuosa
b var. pseudomuyldermansii (Arts, Nordhorn-Richter & A.J.E.Sm.) A.J.E.Sm.
16 P. lescuriana (Sull.) Ochi
17 P. longicolla (Hedw.) Lindb.
18 P. ludwigii (Spreng. ex Schwägr.) Broth.
19 P. lutescens (Limpr.) H.Lindb.
20 P. melanodon (Brid.) A.J.Shaw
21 P. nutans (Hedw.) Lindb.
a subsp. nutans
b subsp. schimperi (Müll.Hal.) Nyholm
22 P. obtusifolia (Vill. ex Brid.) L.F.Koch
23 P. proligera (Kindb.) Lindb. ex Broth.
24 P. scotica Crundw.
25 P. sphagnicola (Bruch & Schimp.) Broth.
26 P. tundrae A.J.Shaw
27 P. vexans (Limpr.) H.Lindb.
28 P. wahlenbergii (F.Weber & D.Mohr)
A.L.Andrews
a var. calcarea (Warnst.) E.F.Warb.
b var. glacialis (Brid.) E.F.Warb.
c var. wahlenbergii
Schizymenium Harv.
1 S. pontevedrense (Luisier) Sérgio Casas, Cros
& Brugués
Mnioideae L.Söderstr. & N.G.Hodgetts394
149 Cinclidium Sw.
1 C. arcticum (Bruch & Schimp.) Schimp.
2 C. latifolium Lindb.
3 C. minutifolium Broth.395
393
4 C. stygium Sw.
5 C. subrotundum Lindb.
150
Cyrtomnium Holmen
1 C. hymenophylloides (Huebener) T.J.Kop.
2 C. hymenophyllum (Bruch & Schimp.)
Holmen
151
Epipterygium Lindb.
1 E. tozeri (Grev.) Lindb.
152
Mnium Hedw.
1 M. blyttii Bruch & Schimp.
2 M. heterophyllum (Hook.) Schwägr.
3 M. hornum Hedw.
4 M. lycopodioides Schwägr.
5 M. marginatum (Dicks.) P.Beauv.
a var. dioicum (H.Müll.) Crundw.
b var. marginatum
6 M. spinosum (Voit) Schwägr.
7 M. spinulosum Bruch & Schimp.
8 M. stellare Hedw.
9 M. thomsonii Schimp.
153
Plagiomnium T.J.Kop. [Orthomnion Wils.]396
section Plagiomnium
1 P. affine (Blandow ex Funck) T.J.Kop.
[Orthomnion affine (Blandow ex
Funck) T.J.Kop. & Yu Sun]
2 P. curvatulum (Lindb.) Schljakov [Orthomnion curvatulum (Lindb.) T.J.Kop. &
Yu Sun]
3 P. cuspidatum (Hedw.) T.J.Kop. [Orthomnion
cuspidatum (Hedw.) T.J.Kop. & Yu
Sun]
4 P. drummondii (Bruch & Schimp.) T.J.Kop.
[Orthomnion drummondii (Bruch &
Schimp.) T.J.Kop. & Yu Sun]
5 P. elatum (Bruch & Schimp.) T.J.Kop. [Orthomnion elatum (Bruch & Schimp.) T.J.Kop.
& Yu Sun]
6 P. ellipticum (Brid.) T.J.Kop. [Orthomnion
ellipticum (Brid.) T.J.Kop. & Yu Sun]
7 P. medium (Bruch & Schimp.) T.J.Kop.
[Orthomnion medium (Bruch &
Schimp.) T.J.Kop. & Yu Sun]
section Undulata (Hedw.) T.J.Kop.
8 P. confertidens (Lindb. & Arnell) T.J.Kop.
[Orthomnion confertidens (Lindb. &
Arnell) T.J.Kop. & Yu Sun]
Pohlia beringiensis was reported from Arctic Russia new to Europe by Ignatov, Afonina and Ignatova (2006).
The subfamily Mnioideae is in general usage but until now it never seems to have been validly published.
Cinclidium minutifolium was reported from Arctic Russia new to Europe by Koponen and Ignatova (2018).
396
The species of Plagiomnium were transferred to Orthomnion by Koponen and Sun (2017), but there is currently a proposal to conserve the name Plagiomnium (Ochyra, Bednarek-Ochyra, et al. 2017). Consequently Plagiomnium is retained.
394
395
JOURNAL OF BRYOLOGY
4
9 P. undulatum (Hedw.) T.J.Kop. [Orthomnion
undulatum (Hedw.) T.J.Kop. & Yu Sun]
a var. madeirense T.J.Kop. & Sérgio
[Orthomnion undulatum var.
madeirense (T.J.Kop. & Sérgio)
T.J.Kop. & Yu Sun]
b var. undulatum
section Rostrata (Hedw.) T.J.Kop.
10 P. rostratum (Schrad.) T.J.Kop. [Orthomnion
rostratum (Schrad.) T.J.Kop. & Yu Sun]
154
Pseudobryum (Kindb.) T.J.Kop.
1 P. cinclidioides (Huebener) T.J.Kop.
155
Rhizomnium (Broth.) T.J.Kop.
1 R. andrewsianum (Steere) T.J.Kop.
2 R. gracile T.J.Kop.
3 R. magnifolium (Horik.) T.J.Kop.
4 R. pseudopunctatum (Bruch & Schimp.) T.J.Kop.
5 R. punctatum (Hedw.) T.J.Kop.
a var. hermanperssonii T.J.Kop.397
b var. punctatum
5
6
7
8
9
10
11
156
Trachycystis Lindb.
1 T. ussuriensis (Maack & Regel) T.J.Kop.
12
Orthotrichales Dixon
Orthotrichaceae Arn.
Orthotrichoideae Broth.
157 Codonoblepharon Schwägr.
1 C. forsteri (Dicks.) Goffinet [Zygodon forsteri
(Dicks.) Mitt.]398
13
14
15
158
397
399
Lewinskya F.Lara, Garilleti & Goffinet
1 L. acuminata (H.Philib.) F.Lara, Garilleti &
Goffinet [Orthotrichum acuminatum
H.Philib.]
2 L. affinis (Schrad. ex Brid.) F.Lara, Garilleti &
Goffinet [Orthotrichum affine Schrad.
ex Brid., Orthotrichum affine var.
bohemicum Plášek & Sawicki]
3 L. breviseta (F.Lara, Garilleti & Mazimpaka)
F.Lara, Garilleti & Goffinet [Orthotrichum speciosum var. brevisetum
F.Lara, Garilleti & Mazimpaka]
16
17
18
159
53
L. elegans (Schwägr. ex Hook. & Grev.) F.Lara,
Garilleti & Goffinet [Orthotrichum
elegans Schwägr. ex Hook. & Grev.]400
L. fastigiata (Bruch ex Brid.) Vigalondo,
F.Lara & Garilleti [Orthotrichum fastigiatum Bruch ex Brid.]401
L. iberica (F.Lara & Mazimpaka) F.Lara, Garilleti & Goffinet [Orthotrichum ibericum
F.Lara & Mazimpaka]
L. iwatsukii (Ignatov) F.Lara, Garilleti &
Goffinet [Orthotrichum iwatsukii
Ignatov, Orthotrichum laevigatum
var. japonicum (Z.Iwats.) Lewinsky,
Orthotrichum macounii subsp. japonicum Z.Iwats.]402
L. laevigata (J.E.Zetterst.) F.Lara, Garilleti &
Goffinet [Orthotrichum laevigatum
J.E.Zetterst.]
L. lamyana F.Lara, Garilleti, Draper &
Mazimpaka403
L. pylaisii (Brid.) F.Lara, Garilleti & Goffinet
[Orthotrichum pylaisii Brid.]
L. rupestris (Schleich. ex Schwägr.) F.Lara,
Garilleti & Goffinet [Orthotrichum
rupestre Schleich. ex Schwägr.]
L. shawii (Wilson) F.Lara, Garilleti & Goffinet
[Orthotrichum shawii Wilson]
L. sordida (Sull. & Lesq.) F.Lara, Garilleti &
Goffinet [Orthotrichum sordidum
Sull. & Lesq.]
L. speciosa (Nees) F.Lara, Garilleti & Goffinet
[Orthotrichum speciosum Nees]
L. striata (Hedw.) F.Lara, Garilleti & Goffinet
[Orthotrichum striatum Hedw.]
L. tortidontia (F.Lara, Garilleti & Mazimpaka)
F.Lara, Garilleti & Goffinet [Orthotrichum tortidontium F.Lara, Garilleti &
Mazimpaka]
L. transcaucasica Eckstein, Garilleti &
F.Lara404
L. vladikavkana (Venturi) F.Lara, Garilleti &
Goffinet [Orthotrichum vladikavkana
Venturi]
Nyholmiella Holmen & E.Warncke405
Rhizomnium punctatum var. hermanperssonii was described from Madeira by Koponen (2017).
Molecular evidence (Goffinet, Shaw, et al. 2004) suggests that Zygodon forsteri is best placed in Codonoblepharon. This is supported by current thinking
(Mazimpaka and Lara 2014).
399
Lewinskya was established as a genus to accommodate the monoicous and phaneroporous taxa of Orthotrichum (Lara, Garilleti, et al. 2016).
400
Vitt and Darigo (1997) established that Lewinskya elegans was distinct from Lewinskya speciosa in North America. It is now known to be widespread in
European Russia (Fedosov and Doroshina 2018), and probably overlooked elsewhere in Europe.
401
Lewinskya fastigiata was reinstated by Vigalondo, Garilleti, et al. (2019), on the basis of integrative taxonomic analyses on the Lewinskya
affinis complex.
402
Lewinskya iwatsukii is an Asian taxon described as a subspecies of Orthotrichum macounii Austin by Iwatuski (1959), raised to species level by Ignatov,
Ivanova, et al. (2001), and later discovered in Europe (Fedosov and Doroshina 2018).
403
Lewinskya lamyana was described from Spain by Lara, Garilleti, et al. (2018).
404
Lewinskya transcaucasica is a recently described species (Eckstein et al. 2018), which has subsequently been found in the European Caucasus by Fedosov,
Doroshina, et al. (2017).
405
The recognition of Nyholmiella at generic level is supported by molecular evidence (Goffinet, Shaw, et al. 2004; Sawicki et al. 2010).
398
54
N. G. HODGETTS ET AL.
1
2
160
406
30 O. shevockii Lewinsky-Haapasaari &
D.H.Norris414
31 O. sibiricum (Grönvall) Warnst. [Orthotrichum
holmenii Lewinsky-Haapasaari]415
32 O. sprucei Mont.
33 O. stellatum Brid.
34 O. stramineum Hornsch. ex Brid.
35 O. tenellum Bruch ex Brid.
36 O. urnigerum Myrin
37 O. vittii F.Lara, Garilleti & Mazimpaka
N. gymnostoma (Bruch ex Brid.) Holmen &
E.Warncke [Orthotrichum gymnostomum Bruch ex Brid.]
N. obtusifolia (Brid.) Holmen & E.Warncke
[Orthotrichum obtusifolium Brid.]
Orthotrichum Hedw.
1 O. alpestre Bruch & Schimp.
2 O. anomalum Hedw.
3 O. bistratosum (Schiffn.) Guerra [Orthotrichum cupulatum var. bistratosum
Schiffn.]406
4 O. callistomum Fisch.-Oost. ex Bruch &
Schimp.
5 O. cambrense Bosanquet & F.Lara407
6 O. casasianum F.Lara, Garilleti & Mazimpaka
7 O. columbicum Mitt. [Orthotrichum consimile auct. eur., non Mitt.]408
8 O. comosum F.Lara, R.Medina & Garilleti409
9 O. consobrinum Cardot410
10 O. crenulatum Mitt.
11 O. cupulatum Brid.
a var. cupulatum
b var. fuscum (Venturi) Boulay
c var. riparium Huebener
12 O. dagestanicum Fedosov & Ignatova411
13 O. dentatum T.Kiebacher & Lüth412
14 O. diaphanum Brid.
15 O. handiense F.Lara, Garilleti & Mazimpaka
16 O. hispanicum F.Lara, Garilleti & Mazimpaka
17 O. macrocephalum F.Lara, Garilleti &
Mazimpaka
18 O. microcarpum De Not.
19 O. moravicum Plášek & Sawicki413
20 O. pallens Bruch ex Brid.
21 O. patens Bruch ex Brid.
22 O. pellucidum Lindb.
23 O. philibertii Venturi
24 O. pulchellum Brunt.
25 O. pumilum Sw. ex anon.
26 O. rivulare Turner
27 O. rogeri Brid.
28 O. scanicum Grönvall
29 O. schimperi Hammar
161
Plenogemma Plášek, Sawicki & Ochyra416
1 P. phyllantha (Brid.) Sawicki, Plášek & Ochyra
[Ulota phyllantha Brid.)
162
Pulvigera Plášek, Sawicki & Ochyra417
1 P. lyellii (Hook. & Taylor) Plášek, Sawicki &
Ochyra [Orthotrichum lyellii Hook. &
Taylor]
163
Ulota D.Mohr
1 U. bruchii Hornsch. ex Brid.
2 U. calvescens Wilson
3 U. coarctata (P.Beauv.) Hammar
4 U. crispa (Hedw.) Brid.418
5 U. crispula Bruch [Ulota crispa var. crispula
(Bruch) Hammar]
6 U. curvifolia (Wahlenb.) Lilj.
7 U. drummondii (Hook. & Grev.) Brid.
8 U. hutchinsiae (Sm.) Hammar
9 U. intermedia Schimp. [Ulota crispa var.
intermedia (Schimp.) Cardot]
10 U. macrospora E.Bauer & Warnst.
11 U. rehmannii Jur.
164
Zygodon Hook. & Taylor
1 Z. catarinoi C.Garcia, F.Lara, Sérgio & Sim-Sim
[Zygodon bistratus Calabrese &
J.Muñoz]419
2 Z. conoideus (Dicks.) Hook. & Taylor
a var. conoideus
b var. lingulatus S.R.Edwards
3 Z. dentatus (Limpr.) Kartt.
4 Z. gracilis Wilson
Orthotrichum cupulatum var. bistratosum was raised to species level by Guerra (1985), although it has been recognised as such only recently (Lara and
Garilleti 2014).
Bosanquet and Lara (2012) described Orthotrichum cambrense from Wales.
408
Orthotrichum columbicum was recognised as a species distinct from Orthotrichum consimile Mitt. by Medina et al. (2012). Orthotrichum consimile s.str. is
confined to the west coast of North America.
409
Orthotrichum comosum was described from southern Europe by Medina et al. (2013).
410
Orthotrichum consobrinum is an Asian species detected in Europe (Spain) by Lara, Mazimpaka, et al. (2009).
411
Orthotrichum dagestanicum was described from the Caucasus by Fedosov and Ignatova (2010).
412
Orthotrichum dentatum was described from the Alps by Kiebacher and Lüth (2017).
413
Orthotrichum moravicum was described from central Europe by Plášek et al. (2009).
414
Orthotrichum shevockii is a disjunct species described from California and recently found in the Canary Islands (Vigalondo, Patiño, et al. 2019).
415
Orthotrichum sibiricum is a north Asian species with one known occurrence in European Arctic Russia (Fedosov et al. 2017b).
416
Sawicki et al. (2017) provide molecular evidence for the treatment of Plášek et al. (2015), segregating Ulota phyllantha in the genus Plenogemma.
417
Sawicki et al. (2017) provide molecular evidence for the treatment of Plášek et al. (2015), segregating Orthotrichum lyellii in the genus Pulvigera.
418
The treatment of the Ulota crispa complex (Ulota crispa s.str., Ulota crispula, Ulota intermedia) follows Caparrós et al. (2016).
419
Zygodon catarinoi was described from Spain, Portugal and Morocco by Garcia et al. (2006), and has subsequently been found elsewhere in the Mediterranean region.
407
JOURNAL OF BRYOLOGY
5
6
7
8
Z. rupestris Schimp. ex Lorentz
Z. sibiricus Ignatov, Ignatova, Z.Iwats. & B.C.Tan
Z. stirtonii Schimp. ex Stirt.
Z. viridissimus (Dicks.) Brid.
Orthodontiales N.E.Bell, A.E.Newton & D.Quandt
Orthodontiaceae Goffinet
165 Leptotheca Schwägr.
1 L. gaudichaudii Schwägr.
166
Orthodontium Schwägr.
1 O. gracile (Wilson) Schwägr. ex Bruch &
Schimp.
2 O. lineare Schwägr.
3 O. pellucens (Hook.) Bruch & Schimp.
Aulacomniales N.E.Bell, A.E.Newton & D.Quandt
Aulacomniaceae Schimp.
167 Aulacomnium Schwägr.
1 A. androgynum (Hedw.) Schwägr.
2 A. palustre (Hedw.) Schwägr.
3 A. turgidum (Wahlenb.) Schwägr.
175
Daltoniaceae Schimp.
170 Achrophyllum Vitt & Crosby
1 A. dentatum (Hook.f. & Wilson) Vitt & Crosby
171
Calyptrochaeta Desv.
1 C. apiculata (Hook.f. & Wilson) Vitt
172
Daltonia Hook. & Taylor
1 D. splachnoides (Sm.) Hook. & Taylor
2 D. lindigiana Hampe [Daltonia stenophylla
Mitt.]420
173
Distichophyllum Dozy & Molk.
1 D. carinatum Dixon & W.E.Nicholson
Hookeriaceae Schimp.
174 Hookeria J.E.Sm.
1 H. lucens (Hedw.) Sm.
Leucomiaceae Broth.
420
Tetrastichium (Mitt.) Cardot
1 T. fontanum (Mitt.) Cardot
2 T. virens (Cardot) S.P.Churchill
Pilotrichaceae Kindb.
Hypnelloideae Broth.
176 Cyclodictyon Mitt.
1 C. laetevirens (Hook. & Taylor) Mitt.
Hypnales W.R.Buck & Vitt
Fontinalaceae Schimp.
177 Dichelyma Myrin
1 D. capillaceum (L. ex Dicks.) Myrin
2 D. falcatum (Hedw.) Myrin
178
Rhizogoniales Goffinet & W.R.Buck
Rhizogoniaceae Broth.
168 Calomnion Hook.f. & Wilson
1 C. complanatum (Hook.f. & Wilson) Lindb.
Hookeriales M.Fleisch. [Hypopterygiales Goffinet]
Hypopterygiaceae Mitt.
169 Hypopterygium Brid.
1 H. tamarisci (Sw.) Brid. ex Müll.Hal.
55
Fontinalis Hedw.
1 F. antipyretica Hedw.
a subsp. antipyretica
b subsp. bryhnii (Limpr.) Podp.
c subsp. gracilis (Lindb.) Kindb.
d subsp. kindbergii (Renauld & Cardot)
Cardot
2 F. dalecarlica Schimp.
3 F. dichelymoides Lindb.
4 F. hypnoides C.Hartm.
a var. duriaei (Schimp.) Kindb.
b var. hypnoides
5 F. squamosa Hedw. [Fontinalis squamosa var.
curnowii Cardot, Fontinalis squamosa
var. dixonii (Cardot) A.J.E.Sm.]421
Plagiotheciaceae M.Fleisch.
179
Herzogiella Broth.
1 H. seligeri (Brid.) Z.Iwats.
2 H. striatella (Brid.) Z.Iwats.
3 H. turfacea (Lindb.) Z.Iwats.
180
Isopterygiopsis Z.Iwats.
1 I. alpicola (Lindb. & Arnell) Hedenäs
2 I. muelleriana (Schimp.) Z.Iwats.
3 I. pulchella (Hedw.) Z.Iwats.
181
Plagiothecium Bruch & Schimp. [Buckiella
Ireland]
1 P. berggrenianum Frisvoll
2 P. cavifolium (Brid.) Z.Iwats.
3 P. curvifolium Schlieph. ex Limpr.
4 P. denticulatum (Hedw.) Schimp.
a var. denticulatum
b var. obtusifolium (Turner) Moore
c var. undulatum R.Ruthe ex Geh.
5 P. laetum Schimp.
6 P. latebricola Schimp.
Majestyk (2011) showed that the correct name for Daltonia stenophylla is Daltonia lindigiana.
We follow Hill et al. (2008) and Guerra (2014) in treating Fontinalis squamosa var. dixonii as synonymous with the type, and var. curnowii is also considered
to have no taxonomic value.
421
56
N. G. HODGETTS ET AL.
7 P. neckeroideum Schimp. [Plagiothecium
noricum Molendo ex Limpr.]422
8 P. nemorale (Mitt.) A.Jaeger
9 P. piliferum (Sw.) Schimp.
10 P. platyphyllum Mönk.
11 P. rossicum Ignatov & Ignatova423
12 P. succulentum (Wilson) Lindb.
13 P. svalbardense Frisvoll
14 P. undulatum (Hedw.) Schimp. [Buckiella
undulata (Hedw.) Ireland]
182
Myurella Bruch & Schimp.
1 M. julacea (Schwägr.) Schimp.
2 M. sibirica (Müll.Hal.) Reimers
3 M. tenerrima (Brid.) Lindb.
183
Ortholimnobium Dixon
1 O. handelii (Broth.) C.Schröck & J.T.Wynns
[Plagiothecium handelii Broth.]424
184
Orthothecium Bruch & Schimp.
1 O. chryseon (Schwägr.) Schimp.
2 O. intricatum (Hartm.) Schimp.
3 O. lapponicum (Schimp.) C.Hartm.
4 O. rufescens (Dicks. ex Brid.) Schimp.
5 O. strictum Lorentz
185
Platydictya Berk.
1 P. jungermannioides (Brid.) H.A.Crum
186
Pseudotaxiphyllum Z.Iwats.
1 P. elegans (Brid.) Z.Iwats.
2 P. laetevirens (Dixon & Luisier ex F.Koppe &
Düll) Hedenäs
Fabroniaceae Schimp.
187
Fabronia Raddi
1 F. altaica Ignatova & Ignatov425
2 F. ciliaris (Brid.) Brid.
3 F. major De Not.426
4 F. pusilla Raddi
Pterigynandraceae Schimp.
188
422
Pterigynandrum Hedw.
1 P. filiforme Hedw. [Pterigynandrum filiforme
var. majus (De Not.) De Not.]427
Habrodontaceae Schimp.
189 Habrodon Schimp.
1 H. perpusillus (De Not.) Lindb.
Climaciaceae Kindb.
190 Climacium F.Weber & D.Mohr
1 C. dendroides (Hedw.) F.Weber & D.Mohr
Myriniaceae Schimp.
191 Myrinia Schimp.
1 M. pulvinata (Wahlenb.) Schimp.
Amblystegiaceae G.Roth
Cratoneuroideae Vanderp., Hedenäs,
A.J.Shaw
192 Cratoneuron (Sull.) Spruce
1 C. curvicaule (Jur.) G.Roth
2 C. filicinum (Hedw.) Spruce
193
C.J.Cox
&
Palustriella Ochyra
1 P. commutata (Hedw.) Ochyra
2 P. decipiens (De Not.) Ochyra
3 P. falcata (Brid.) Hedenäs [Palustriella commutata var. sulcata (Lindb.) Ochyra,
Palustriella pluristratosa M.Stech & J.P.Frahm]428
Amblystegioideae Broth.
194 Amblystegium Schimp.
1 A. serpens (Hedw.) Schimp. [Amblystegium
serpens var. salinum Carrington]
195
Anacamptodon Brid.
1 A. splachnoides (Froel. ex Brid.) Brid.
196
Arvernella Hugonnot & Hedenäs429
1 A. microclada Hugonnot & Hedenäs
197
Campyliadelphus (Kindb.) R.S.Chopra
1 C. chrysophyllus (Brid.) R.S.Chopra
2 C. elodes (Lindb.) Kanda
198
Campylium (Kindb.) R.S.Chopra
1 C. bambergeri (Schimp.) Hedenäs, Schlesak &
D.Quandt
[Hypnum
bambergeri
430
Schimp.]
2 C. laxifolium Engelmark & Hedenäs
The status of Plagiothecium noricum Molendo ex. Limpr., and its relation to Plagiothecium neckeroideum, still needs clarification.
Plagiothecium rossicum was descibred from Russia by Ignatova, Fedorova, et al. (2019).
Plagiothecium handelii was recorded new to Europe, and its position in Ortholimnobium confirmed, by Wynns and Schröck (2018).
425
Fabronia altaica was described from Russia by Ignatova, Kuznetsova, et al. (2017). In Europe it is restricted to the Caucasus.
426
Fabronia major was resurrected in the course of a study on the genus Fabronia in Russia by Ignatova, Kuznetsova, et al. (2017).
427
According to Hill et al. (2008), ”Smith (2004) retains Pterigynandrum filiforme var. majus but expresses severe doubts as to whether it is distinct. It is treated
here as a synonym of var. filiforme.” We follow this treatment.
428
The synonymy of Palustriella pluristratosa with Palustriella falcata follows Hedenäs (2010). The taxonomic value of Palustriella commutata var. sulcata
should be studied in more detail. It is a characteristic plant of high altitude limestone in the Alps.
429
Arvernella is a new genus, described by Hugonnot and Hedenäs (2015) to accommodate Arvernella microclada, a new species found in the Auvergne,
France. Its placement in the Amblystegioideae is supported by recent analysis by Kučera, Kuznetsova, et al. (2019).
430
Molecular evidence places Hypnum bambergeri in Campylium (Schlesak et al. 2018).
423
424
JOURNAL OF BRYOLOGY
3
4
5
199
200
Campylophyllum (Schimp.) M.Fleisch.
1 C. halleri (Hedw.) M.Fleisch.
2 C. montanum (Lindb.) B.H.Allen [Hygrohypnum montanum (Lindb.) Broth., Platyhypnum
montanum
(Lindb.)
433
Ochyra]
201
Conardia H.Rob.434
1 C. compacta (Drumm. ex Müll.Hal.) H.Rob.
202
Drepanium (Schimp.) C.E.O.Jensen435
1 D. fastigiatum (Hampe) C.E.O.Jensen
[Hypnum recurvatum (Lindb. &
Arnell) Kindb., Drepanium recurvatum
(Lindb. & Arnell) G.Roth]
203
431
5
D. capillifolius (Warnst.) Warnst. [Drepanocladus longifolius auct. eur., non
(Mitt.) Paris]437
6 D. lycopodioides (Brid.) Warnst. [Pseudocalliergon lycopodioides (Brid.) Hedenäs]
7 D. polygamus (Schimp.) Hedenäs438
8 D. sendtneri (Schimp. ex H.Müll.) Warnst.
9 D. sordidus (Müll.Hal.) Hedenäs
10 D. trifarius (F.Weber & D.Mohr) Broth. ex
Paris [Pseudocalliergon trifarium
(F.Weber & D.Mohr) Loeske]
11 D. turgescens (T.Jensen) Broth. [Pseudocalliergon turgescens (T.Jensen) Loeske]
C. longicuspis (Lindb. & Arnell) Hedenäs
C. protensum (Brid.) Kindb.431
C. stellatum (Hedw.) Lange & C.E.O.Jensen
Campylophyllopsis W.R.Buck [Campylidium
(Kindb.) Ochyra, nom. illeg.]432
1 C. calcarea (Crundw. & Nyholm) Ochyra
[Campylidium calcareum (Crundw. &
Nyholm) Ochyra, Campylophyllum
calcareum (Crundw. & Nyholm)
Hedenäs]
2 C. sommerfeltii (Myrin) Ochyra [Campylidium
sommerfeltii (Myrin) Ochyra, Campylophyllum
sommerfeltii
(Myrin)
Hedenäs]
Drepanocladus (Müll.Hal.) G.Roth [Pseudocalliergon (Limpr.) Loeske]436
1 D. aduncus (Hedw.) Warnst.
2 D. angustifolius (Hedenäs) Hedenäs &
C.Rosborg [Pseudocalliergon angustifolium Hedenäs]
3 D. arcticus (R.S.Williams) Hedenäs
4 D. brevifolius (Lindb.) Warnst. [Pseudocalliergon
brevifolium
(Lindb.)
Hedenäs]
57
204
Hygroamblystegium Loeske439
1 H. fluviatile (Hedw.) Loeske
2 H. humile (P.Beauv.) Vanderp., Goffinet &
Hedenäs [Hygroamblystegium varium
var. humile Vanderp. & Hedenäs]440
3 H. tenax (Hedw.) Jenn.
4 H. varium (Hedw.) Mönk. [Amblystegium
varium (Hedw.) Lindb.]
205
Hygrohypnum Lindb.
1 H. luridum (Hedw.) Jenn.
2 H. styriacum (Limpr.) Broth.
206
Leptodictyum (Schimp.) Warnst.
1 L. riparium (Hedw.) Warnst.
207
Microhypnum Jan Kučera & Ignatov
1 M. sauteri (Schimp.) Jan Kučera & Ignatov
[Anacamptodon sauteri (Schimp.)
Hedenäs, Schlesak & D.Quandt,
Hypnum sauteri Schimp.]441
208
Platyhypnum Loeske [Ochyraea Váňa]442
1 P. alpestre (Hedw.) Ochyra [Hygrohypnum
alpestre (Hedw.) Loeske]
2 P. alpinum (Lindb.) Loeske [Hygrohypnum
alpinum (Lindb.) Loeske]
Campylium protensum and Campylium stellatum are distinct in some parts of their range and overlap in others. They are retained as separate species for
the present, although there is a strong case for Campylium protensum to be reduced to a variety of Campylium stellatum, as some authors have done.
The separation of Campylidium from Campylophyllum is supported by molecular studies (Gardiner et al. 2005; Ignatov, Gardiner, et al. 2007). However,
Campylidium is an illegitimate name and must be changed to Campylophyllopsis (Goffinet, Buck, et al. 2009).
433
Ochyra (2013) placed Hygrohypnum montanum in Platyhypnum. Allen (2014) transferred it to Campylophyllum.
434
Although here retained in the Amblystegiaceae, Conardia does not appear to belong here, and may require a family of its own (see, for example, Vanderpoorten et al. 2002).
435
Recent molecular studies place Hypnum recurvatum in the genus Drepanium (Schlesak et al. 2018).
436
This treatment of Drepanocladus follows Hedenäs and Rosborg (2008).
437
Drepanocladus longifolius (Mitt.) Paris is an exclusively Southern Hemisphere species; Saługa et al. (2018) showed that European records of this species
should be referred to Drepanocladus capillifolius.
438
Campylium decipiens (Warnst.) Walsemann is a central European plant recognised as a distinct species by Frahm and Walsemann (1973) and Meinunger
and Schröder (2007), but otherwise largely disregarded. It may be a form of Drepanocladus polygamus, but it requires further study.
439
Although Hygroamblystegium fluviatile, Hygroamblystegium humile and Hygroamblystegium tenax have been synonymised with Hygroamblystegium
varium (Vanderpoorten 2004), we follow Hill et al. (2006) in retaining these species as distinct pending further work.
440
Hygroamblystegium humile was recognised at varietal level by Vanderpoorten and Hedenäs (2009), while Hygroamblystegium fluviatile and Hygroamblystegium tenax continued to be regarded as synonyms of Amblystegium varium.
441
On molecular evidence, Schlesak et al. (2018) placed Hypnum sauteri in Anacamptodon but Kučera, Kuznetsova, et al. (2019) provided further evidence
placing it in the new genus Microhypnum.
442
We follow the treatment of Ochyra (2013) in transferring several Hygrohypnum species to Platyhypnum.
432
58
N. G. HODGETTS ET AL.
3
4
5
6
7
8
P. cochlearifolium (Venturi) Ochyra [Hygrohypnum cochlearifolium (Venturi) Broth.]
P. duriusculum (De Not.) Ochyra [Hygrohypnum
duriusculum
(De
Not.)
D.W.Jamieson]
P. molle (Dicks. ex Hedw.) Loeske [Hygrohypnum molle (Dicks. ex Hedw.) Loeske]
P. norvegicum (Schimp.) Ochyra [Hygrohypnum norvegicum (Schimp.) J.J.Amann]
P. smithii (Sw.) Ochyra [Hygrohypnum smithii
(Sw.) Broth.]
P. tatrense (Váňa) Hedenäs & Ignatov [Ochyraea tatrensis Váňa]443
209
Pseudoamblystegium Vanderp. & Hedenäs444
1 P. subtile (Hedw.) Vanderp. & Hedenäs [Amblystegium subtile (Hedw.) Schimp.]
210
Pseudocampylium Vanderp. & Hedenäs445
1 P. radicale (P.Beauv.) Vanderp. & Hedenäs
[Amblystegium radicale (P.Beauv.)
Schimp.]
211
Serpoleskea (Limpr.) Loeske
1 S. confervoides (Brid.) Schimp. [Amblystegium
confervoides (Brid.) Schimp.]446
212
Tomentypnum Loeske
1 T. nitens (Hedw.) Loeske
3 S. sarmentosum (Wahlenb.) Tuom. & T.J.Kop.
[Warnstorfia sarmentosa (Wahlenb.)
Hedenäs]
4 S. trichophyllum (Warnst.) Hedenäs [Warnstorfia trichophylla (Warnst.) Tuom. &
T.J.Kop.]
5 S. tundrae (Arnell) Hedenäs [Warnstorfia
tundrae (Arnell) Loeske]
216
Straminergon Hedenäs
1 S. stramineum (Dicks. ex Brid.) Hedenäs
217
Warnstorfia Loeske
1 W. fluitans (Hedw.) Loeske
2 W. pseudostraminea (Müll.Hal.) Tuom. &
T.J.Kop.
Scorpidiaceae Ignatov & Ignatova
218 Hamatocaulis Hedenäs
1 H. lapponicus (Norrl.) Hedenäs
2 H. vernicosus (Mitt.) Hedenäs448
219
Hygrohypnella Ignatov & Ignatova449
1 H. ochracea (Turner ex Wilson) Ignatov &
Ignatova [Hygrohypnum ochraceum
(Turner ex Wilson) Loeske]
2 H. polaris (Lindb.) Ignatov & Ignatova [Hygrohypnum polare (Lindb.) Loeske]
220
Sanionia Loeske
1 S. nivalis Hedenäs [Sanionia georgicouncinata auct. eur., non (Müll.Hal.)
Ochyra & Hedenäs]450
2 S. orthothecioides (Lindb.) Loeske
3 S. uncinata (Hedw.) Loeske
221
Scorpidium (Schimp.) Limpr.
1 S. cossonii (Schimp.) Hedenäs
2 S. revolvens (Sw. ex anon.) Rubers
3 S. scorpioides (Hedw.) Limpr.
Calliergonaceae Vanderp., Hedenäs, C.J.Cox & A.J.Shaw
213
Calliergon (Sull.) Kindb.
1 C. cordifolium (Hedw.) Kindb.
2 C. giganteum (Schimp.) Kindb.
3 C. megalophyllum Mikut.
4 C. richardsonii (Mitt.) Kindb.
214
Loeskypnum H.K.G.Paul
1 L. badium (Hartm.) H.K.G.Paul
215
Sarmentypnum Tuom. & T.J.Kop.447
1 S. exannulatum (Schimp.) Hedenäs [Warnstorfia exannulata (Schimp.) Loeske]
2 S. procerum (Renauld & Arnell) Hedenäs
[Warnstorfia procera (Renauld &
Arnell) Tuom.]
443
Leskeaceae Schimp.
222 Claopodium (Lesq. & James) Renauld & Cardot
1 C. rostratum (Hedw.) Ignatov [Anomodon rostratus (Hedw.) Schimp.]451
2 C. whippleanum (Sull.) Renauld & Cardot
Ochyraea tatrensis may be nothing more than an extreme form of Platyhypnum smithii, but for the present it is retained as a species and transferred to the
genus Platyhypnum until further work is done.
The genus Pseudoamblystegium was established by Vanderpoorten and Hedenäs (2009) to accommodate Amblystegium subtile.
445
The genus Pseudocampylium was established by Vanderpoorten and Hedenäs (2009) to accommodate Amblystegium radicale.
446
The placement of Amblystegium confervoides in Serpoleskea (Vanderpoorten et al. 2002) was confirmed by Vanderpoorten and Hedenäs (2009).
447
The treatment of Sarmentypnum follows Hedenäs (2006).
448
Hamatocaulis vernicosus comprises two well-researched cryptic species in Europe. Careful morphological evaluation has failed to find any morphological
difference between the two (Manukjanová et al. 2019).
449
Hygrohypnum ochraceum and Hygrohypnum polare are placed in Hygrohypnella, following Ignatov and Ignatova (2004).
450
Molecular studies showed European Sanionia nivalis to be distinct from the Southern Hemisphere Sanionia georgicouncinata (Müll.Hal.) Ochyra & Hedenäs
(Hedenäs 2012).
451
Molecular studies place Anomodon rostratus in Claopodium (Ignatov, Afonina, Ignatova 2006).
444
JOURNAL OF BRYOLOGY
223
Leskea Hedw.
1 L. polycarpa Hedw.
224
Lindbergia Kindb.452
1 L. dagestanica Ignatova & Ignatov
2 L. grandiretis (Lindb. ex Broth.) Ignatov &
Ignatova [Lindbergia brachyptera
auct. eur., non (Mitt.) Kindb.]
225
Pseudoleskeopsis Broth.453
1 P. artariae (Thér.) Thér. [Pseudoleskea artariae
Thér.]
Pseudoleskeaceae Schimp.
226 Lescuraea Bruch & Schimp. [Pseudoleskea
Schimp., Ptychodium Schimp.]454
1 L. incurvata (Hedw.) E.Lawton [Pseudoleskea
incurvata (Hedw.) Loeske]
2 L. mutabilis (Brid.) Lindb. ex I.Hagen
3 L. patens Lindb. [Pseudoleskea patens (Lindb.)
Kindb.
4 L. plicata (Schleich. ex F.Weber & D.Mohr)
Broth.
[Ptychodium
plicatum
(Schleich. ex F.Weber & D.Mohr)
Schimp.]
5 L. radicosa (Mitt.) Mönk. [Pseudoleskea radicosa (Mitt.) Macoun & Kindb.]
6 L. saviana (De Not.) E.Lawton [Pseudoleskea
saviana (De Not.) Latzel]
7 L. saxicola (Schimp.) Molendo
8 L. secunda Arnell
Pseudoleskeellaceae Ignatov & Ignatova
227 Pseudoleskeella Kindb.
1 P. catenulata (Brid. ex Schrad.) Kindb.
2 P. nervosa (Brid.) Nyholm
3 P. papillosa (Lindb.) Kindb.
4 P. rupestris (Berggr.) Hedenäs & L.Söderstr.
5 P. tectorum (Funck ex Brid.) Kindb. ex Broth.
Thuidiaceae Schimp.
228 Abietinella Müll.Hal.
1 A. abietina (Hedw.) M.Fleisch.
a var. abietina
b var. hystricosa (Mitt.) Sakurai
229
452
Haplocladium (Müll.Hal.) Müll.Hal.
1 H. angustifolium (Hampe & Müll.Hal.) Broth.
59
2 H. microphyllum (Hedw.) Broth.
3 H. virginianum (Brid.) Broth.
230
Helodium (Müll.Hal.) Müll.Hal.455
1 H. blandowii (F.Weber & D.Mohr) Warnst.
[Elodium blandowii (F.Weber &
D.Mohr) Eckel]
231
Pelekium Mitt.
1 P. atlanticum (Hedenäs) Hedenäs
2 P. minutulum (Hedw.) Touw
232
Thuidiopsis (Broth.) M.Fleisch.
1 T. sparsa (Hook.f. & Wilson) Broth.
233
Thuidium Bruch & Schimp.
1 T. assimile (Mitt.) A.Jaeger
2 T. delicatulum (Hedw.) Schimp.
3 T. recognitum (Hedw.) Lindb.
4 T. tamariscinum (Hedw.) Schimp.
Brachytheciaceae Schimp.
Eurhynchioideae Milde
234
Eurhynchium Bruch & Schimp.
1 E. angustirete (Broth.) T.J.Kop.
2 E. striatum (Hedw.) Schimp.
235
Palamocladium M.Fleisch.
1 P. euchloron (Müll.Hal.) Wijk & Margad.
236
Plasteurhynchium M.Fleisch.
1 P. meridionale (Schimp.) M.Fleisch.
2 P. striatulum (Spruce) M.Fleisch.
237
Pseudoscleropodium (Limpr.) M.Fleisch.
1 P. purum (Hedw.) M.Fleisch.
238
Rhynchostegium Bruch & Schimp. [Platyhypnidium M.Fleisch.]456
1 R. alopecuroides (Brid.) A.J.E.Sm. [Platyhypnidium lusitanicum (Schimp.) Ochyra &
Bedn.-Ochyra,
Platyhypnidium
mutatum Ochyra & Vanderp.]457
2 R. confertum (Dicks.) Schimp.
3 R. confusum K.Cezón, J.Muñoz, Hedenäs &
Huttunen458
The treatment of Lindbergia follows Ignatova, Ignatov, et al. (2010).
The type of Pseudoleskea is Pseudoleskea atrovirens (=Pseudoleskea incurvata), now in Lescuraea, so Pseudoleskea artariae cannot be the sole representative
of the genus. Therefore we return this taxon to the genus Pseudoleskeopsis (placed by Frey and Stech 2009 in Leskeaceae) pending further studies.
454
Molecular studies suggest that Pseudoleskea and Ptychodium should be included in Lescuraea (Gardiner et al. 2005; Ignatov, Gardiner, et al. 2007).
455
Eckel (2012) proposed that Elodium (Sull.) Austin take the place of Helodium, on the basis of precedence. However, preliminary research by M. Ignatov
indicates that North American and European material is not necessarily congeneric. Globally, there are (traditionally) three species commonly placed in the
genus Helodium, but they are probably unrelated. As the name Helodium is in common usage, it is retained pending further studies.
456
Molecular studies have shown that the genus Platyhypnidium is not sustainable, with all the European species correctly placed in Rhynchostegium (Huttunen and Ignatov 2010).
457
The generic placement of Rhynchostegium alopecuroides was confirmed using molecular techniques by Huttunen and Ignatov (2010). Molecular studies
have shown that Platyhypnidium mutatum is deeply nested within Rhynchostegium alopecuroides, although a formal synonymy was not made (Hutsemékers et al. 2012).
458
Rhynchostegium confusum was described from the Iberian Peninsula by Cezón et al. (2010).
453
60
N. G. HODGETTS ET AL.
4
5
6
7
8
239
R. megapolitanum (Blandow ex F.Weber &
D.Mohr) Schimp.
R. murale (Hedw.) Schimp. [Rhynchostegium
arcticum
(I.Hagen)
Ignatov
&
459
Huttunen]
R. riparioides (Hedw.) Cardot [Platyhypnidium
grolleanum Ochyra & Bedn.-Ochyra,
Platyhypnidium torrenticola (Ochyra,
C.Schmidt & Bültmann) Ochyra &
Bedn.-Ochyra]460
R. rotundifolium (Scop. ex Brid.) Schimp.
R. strongylense (Bott.) W.R.Buck & Privitera
Scorpiurium Schimp.
1 S. circinatum (Bruch) M.Fleisch. & Loeske
2 S. deflexifolium (Solms) M.Fleisch. & Loeske
3 S. sendtneri (Schimp.) M.Fleisch.
Helicodontoideae M.Fleisch.
240 Cirriphyllum Grout
1 C. crassinervium (Taylor) Loeske & M.Fleisch.
2 C. piliferum (Hedw.) Grout
241
Clasmatodon Hook.f. & Wilson
1 C. parvulus (Hampe) Sull.461
242
Hedenasiastrum Ignatov & Vanderp.462
1 H. percurrens (Hedenäs) Ignatov &
Vanderp. [Brachythecium percurrens
Hedenäs]
243
Helicodontium Schwägr.
1 H. capillare (Hedw.) A.Jaeger
244
Microeurhynchium Ignatov & Vanderp.463
1 M. pumilum (Wilson) Ignatov & Vanderp.
[Oxyrrhynchium pumilum (Wilson)
Loeske]
245
Nobregaea Hedenäs
1 N. latinervis Hedenäs
459
246
Oxyrrhynchium (Schimp.) Warnst.
1 O. hians (Hedw.) Loeske
2 O. schleicheri (R.Hedw.) Röll
3 O. speciosum (Brid.) Warnst.
247
Pseudorhynchostegiella Ignatov & Vanderp.464
1 P. duriaei (Mont.) Ignatov & Vanderp.
[Rhynchostegiella durieui (Mont.)
P.Allorge & Perss.]
248
Rhynchostegiella (Schimp.) Limpr.
1 R. azorica Hedenäs & Vanderp.465
2 R. bourgaeana (Mitt.) Broth.
3 R. curviseta (Brid.) Limpr.
4 R. litorea (De Not.) Limpr. [Rhynchostegiella
tenella var. meridionalis (Boulay)
Zodda]466
5 R. pseudolitorea Hedenäs & J.Patiño467
6 R. tenella (Dicks.) Limpr.
7 R. teneriffae (Mont.) Dirkse & Bouman
[Rhynchostegiella jacquinii (Garov.)
Limpr., Rhynchostegiella macilenta
(Renauld & Cardot) Cardot, Rhynchostegiella teesdalei (Schimp.) Limpr.]468
8 R. trichophylla Dirkse & Bouman
9 R. tubulosa Hedenäs & J.Patiño469
Brachythecioideae Lotsy
249 Brachytheciastrum Ignatov & Huttunen
1 B. collinum (Schleich. ex Müll.Hal.) Ignatov &
Huttunen [Brachytheciastrum fendleri
auct. eur., non (Sull.) Ochyra &
Żarnowiec]470
2 B. dieckei (Röll) Ignatov & Huttunen
3 B. olympicum (Jur.) Vanderp. et al.
4 B. salicinum (Schimp.) J.D.Orgaz, M.J.Cano &
J.Guerra [Brachytheciastrum velutinum var. salicinum (Schimp.) Ochyra
& Żarnowiec]471
5 B. trachypodium (Brid.) Ignatov & Huttunen
6 B. velutinum (Hedw.) Ignatov & Huttunen
[Brachytheciastrum vanekii (Šmarda)
Recent molecular work by M. Ignatov suggests that Rhynchostegium arcticum should be treated as a synonym of Rhynchostegium murale.
Platyhypnidium grolleanum was regarded as a rare mutation of Rhynchostegium riparioides by Kučera, Váňa, et al. (2012). Molecular studies have shown
that Platyhypnidium torrenticola is deeply nested within Rhynchostegium riparioides, although a formal synonymy was not made (Hutsemékers et al. 2012).
461
Clasmatodon parvulus is re-admitted into the European flora following the revision of an old herbarium specimen (collected in Germany in 1851) by Müller
(2007). It has not been refound more recently.
462
Hedenasiastrum was described on molecular grounds (Aigoin et al. 2009).
463
Microeurhynchium was described on molecular grounds (Aigoin et al. 2009).
464
Pseudorhynchostegiella was described on molecular grounds (Aigoin et al. 2009).
465
Rhynchostegiella azorica is an Azorean endemic described by Vanderpoorten et al. (2015).
466
The synonymy of Rhynchostegiella tenella var. meridionalis with Rhynchostegiella litorea follows Guerra, Ríos, et al. (2014) and Patiño et al. (2017).
467
Rhynchostegiella pseudolitorea is a Macaronesian endemic described by Patiño et al. (2017).
468
Rhynchostegiella macilenta was synonymised with R. teneriffae by Patiño et al. (2017). The difficulties with Rhynchostegiella jacquinii and Rhynchostegiella
teesdalei were discussed by Patiño et al. (2017), who listed them as doubtfully synonymous with Rhynchostegiella teneriffae. We continue to treat them as
synonyms pending further work.
469
Rhynchostegiella tubulosa is an eastern Mediterranean species described by Patiño et al. (2017), and subsequently also found in Portugal (Ellis, Amélio,
et al. 2019).
470
All European records of Brachytheciastrum fendleri are Brachytheciastrum collinum (Orgaz et al. 2013). Brachytheciastrum fendleri (Sull.) Ochyra & Żarnowiec
is a North American species.
471
Brachytheciastrum salicinum is reinstateded as a species, as it is significantly different from Brachytheciastrum velutinum morphologically, molecularly and
ecologically (D. Orgaz pers. comm. 2018).
460
JOURNAL OF BRYOLOGY
Ochyra & Żarnowiec, Brachytheciastrum velutinum var. vagans (Milde)
Ochyra & Żarnowiec]472
250
472
Brachythecium Schimp. [Bryhnia Kaurin]
1 B. albicans (Hedw.) Schimp.
2 B. buchananii (Hook.) A.Jaeger473
3 B. campestre (Müll.Hal.) Schimp.
4 B. capillaceum (F.Weber & D.Mohr) Giacom.
[Brachythecium
rotaeanum
De
Not.]474
5 B. cirrosum (Schwägr.) Schimp.
6 B. erythrorrhizon Schimp. [Brachythecium
erythrorrhizon
subsp.
asiaticum
Ignatov, Brachythecium erythrorrhizon
subsp. erythrorrhizon var. thedenii
(Schimp.) Lindb.]475
7 B. funkii Schimp.476
8 B. geheebii Milde
9 B. glareosum (Bruch ex Spruce) Schimp.
10 B. japygum (Głow.) Köckinger & Jan
Kučera477
11 B. laetum (Brid.) Schimp.
12 B. mildeanum (Schimp.) Schimp.
13 B. novae-angliae (Sull. & Lesq.) A.Jaeger
[Brachythecium scabridum (Lindb.)
M.Li & Y.F.Wang, Bryhnia scabrida
(Lindb.) Kaurin; Bryhnia novaeangliae (Sull. & Lesq.) Grout]478
14 B. rivulare Schimp.
15 B. rutabulum (Hedw.) Schimp.
a var. atlanticum Hedenäs
b var. rutabulum
16 B. salebrosum (Hoffm. ex F.Weber &
D.Mohr) Schimp.
17 B. tauriscorum Molendo [Brachythecium
coruscum I.Hagen]479
18 B. tenuicaule (Spruce) Kindb. [Rhynchostegiella tenuicaulis (Spruce) Kartt.]480
61
19 B. tommasinii (Sendtn. ex Boulay) Ignatov &
Huttunen
20 B. turgidum (Hartm.) Kindb.
21 B. udum I.Hagen [Brachythecium mildeanum
var. udum (I.Hagen) Mönk.]481
251
Eurhynchiastrum Ignatov & Huttunen
1 E. diversifolium (Schimp.) J.Guerra [Eurhynchiastrum pulchellum var. diversifolium
(Schimp.) Ochyra & Żarnowiec]482
2 E. pulchellum (Hedw.) Ignatov & Huttunen
[Eurhynchiastrum
pulchellum
var.
praecox (Hedw.) Ochyra & Żarnowiec]483
252
Homalothecium Schimp.
1 H. aureum (Spruce) H.Rob.
2 H. lutescens (Hedw.) H.Rob.
a var. fallax (H.Philib. ex Schimp.) Düll
b var. lutescens
3 H. mandonii (Mitt.)Geh.484
4 H. meridionale (M.Fleisch. & Warnst.)
Hedenäs485
5 H. philippeanum (Spruce) Schimp.
6 H. sericeum (Hedw.) Schimp.
253
Kindbergia Ochyra
1 K. praelonga (Hedw.) Ochyra
254
Myuroclada Besch.
1 M. longiramea (Müll.Hal.) M.Li, Y.-F.Wang,
Ignatov & Huttunen486
2 M. maximowiczii (G.G.Borshch.) Steere &
W.B.Schofield
255
Sciuro-hypnum (Hampe) Hampe
1 S. curtum (Lindb.) Ignatov487
2 S. dovrense (Limpr.) Draper & Hedenäs488
3 S. flotowianum (Sendtn.) Ignatov & Huttunen
4 S. glaciale (Schimp.) Ignatov & Huttunen
5 S. latifolium (Kindb.) Ignatov & Huttunen
Brachytheciastrum vanekii and Brachytheciastrum velutinum var. vagans (the latter known only from Poland) are synonymised with Brachytheciastrum velutinum, as they do not differ either morphologically or molecularly (J. Kučera and D. Orgaz pers. comm. 2018).
Brachythecium buchananii is an essentially Asiatic species that also occurs on the European side of the Ural mountains (Ignatov and Milyutina 2010).
474
Brachythecium capillaceum is most likely the earlier name for Brachythecium rotaeanum, although additional study is needed to confirm this.
475
In reference to Brachythecium erythrorrhizon subsp. asiaticum, Ignatov and Milyutina (2010) observed, ” … too broad variation in plant size in Eurasia and
North America, making [it] very difficult to segregate this subspecies.” Neither Brachythecium erythrorrhizon subsp. asiaticum nor subsp. erythrorrhizon var.
thedenii are now considered worthy of recognition at any level.
476
Brachythecium funkii was segregated from Brachythecium cirrosum on the basis of molecular evidence (Köckinger and Kučera 2016).
477
Brachythecium japygum was segregated from Brachythecium cirrosum on the basis of molecular evidence (Köckinger and Kučera 2016).
478
The Eurasian Brachythecium scabridum was synonymised with the North American Brachythecium novae-angliae by Huttunen, Kuznetsova, et al. (2015), as
no significant differences could be found.
479
Brachythecium tauriscorum is an older name for B. coruscum (Hedenäs 2017b).
480
Molecular work showed Brachythecium tenuicaule to be closely related to Brachythecium tommasinii (Köckinger and Kučera 2016).
481
Brachythecium udum was treated as a variety of Brachythecium mildeanum (Podpera 1954; Hill et al. 2006), but Ignatov and Milyutina (2010) found it to be
more closely related to Brachythecium turgidum. Sporophytes and male gametangia remain unknown, and it needs further study to determine whether it is
a good species or an extreme phenotype in the Brachythecium salebrosum-Brachythecium turgidum complex growing in wet northern environments.
482
Eurhynchiastrum diversifolium was raised to species level by Guerra (2016).
483
The treatment of Eurhynchiastrum pulchellum follows Guerra (2016).
484
Homalothecium mandonii was segregated from H. sericeum on the basis of molecular and morphological evidence (Hedenäs, Désamoré, et al. 2014).
485
Homalothecium meridionale was segregated from H. sericeum on the basis of molecular and morphological evidence (Hedenäs, Désamoré, et al. 2014).
486
The status of Myuroclada longiramea in Europe was clarified by Ignatov, Huttunen, et al. (2015).
487
Sciuro-hypnum curtum was restored from synonymy with Sciuro-hypnum oedipodium by Ignatov and Milyutina (2007). It is a widespread species in Europe,
whereas Sciuro-hypnum oedipodium, which is primarily a western North American species, is very rare, with just a few records from eastern Europe.
488
Sciuro-hypnum dovrense was removed from synonymy with Brachythecium glaciale by Draper and Hedenäs (2009).
473
62
N. G. HODGETTS ET AL.
6
7
8
9
10
11
12
256
S. oedipodium (Mitt.) Ignatov & Huttunen
S. ornellanum (Molendo) Ignatov & Huttunen
S. plumosum (Hedw.) Ignatov & Huttunen
S. populeum (Hedw.) Ignatov & Huttunen
S. reflexum (Starke) Ignatov & Huttunen
S. starkei (Brid.) Ignatov & Huttunen
S. tromsoeense (Kaurin & Arnell) Draper &
Hedenäs489
Scleropodium Bruch & Schimp.
1 S. cespitans (Wilson ex Müll.Hal.) L.F.Koch
2 S. touretii (Brid.) L.F.Koch
Taxiphyllaceae Ignatov496
259 Taxiphyllum M.Fleisch.
1 T. densifolium (Lindb. ex Broth.) Reimers
2 T. wissgrillii (Garov.) Wijk & Margad.
Pylaisiadelphaceae Goffinet & W.R.Buck
260 Brotherella M.Fleisch.
1 B. lorentziana (Molendo ex Lorentz)
Loeske ex M.Fleisch. [Brotherella
henonii auct. eur. non (Duby)
M.Fleisch.]497
261
Heterophyllium (Schimp.) Kindb.
1 H. nemorosum (W.D.J.Koch ex Brid.) Kindb.
[Heterophyllium
affine
(Hook.)
M.Fleisch., nom. illeg.]498
262
Isopterygium Mitt.
1 I. tenerum (Sw.) Mitt.
263
Platygyrium Bruch & Schimp.
1 P. repens (Brid.) Schimp.
Hypnaceae Schimp.
257
Hypnum Hedw.490
1 H. andoi A.J.E.Sm.
2 H. cupressiforme Hedw.
a var. cupressiforme
b var. filiforme Brid. 491
c var. heseleri (Ando & Higuchi) M.O.Hill
d var. lacunosum Brid.
e var. subjulaceum Molendo [Hypnum subcomplanatum Hedenäs, Schlesak
& D.Quandt, nom. illeg., Hypnum
subjulaceum (Molendo) Hedenäs,
Schlesak & D.Quandt]492
3 H. jutlandicum Holmen & E.Warncke
4 H. resupinatum Taylor [Hypnum cupressiforme
var. resupinatum (Taylor) Schimp.]493
5 H. uncinulatum Jur.
Callicladiaceae Jan Kučera & Ignatov494
258
489
Callicladium H.A.Crum
1 C. haldanianum (Grev.) H.A.Crum
2 C. imponens (Hedw.) Hedenäs, Schlesak &
D.Quandt [Hypnum imponens Hedw.]495
Jocheniaceae Jan Kučera & Ignatov499
264 Jochenia Hedenäs, Schlesak & D.Quandt500
1 J. pallescens (Hedw.) Hedenäs, Schlesak &
D.Quandt
[Hypnum
pallescens
(Hedw.) P.Beauv., Hypnum reptile
Michx., Hypnum pallescens var.
reptile (Michx.) Husn.]
2 J. protuberans (Brid.) Jan Kučera & Ignatov
[Hypnum pallescens var. protuberans
(Brid.) Austin, Hypnum protuberans
Brid.]501
Stereodontaceae Hedenäs, Schlesak & D.Quandt502
265 Stereodon (Brid.) Mitt. [Breidleria Loeske]503
Sciuro-hypnum tromsoeense was removed from synonymy with Brachythecium starkei by Draper and Hedenäs (2008).
The treatment of Hypnum follows Schlesak et al. (2018), apart from Hypnum cupressiforme var. subjulaceum (see footnote below).
Hypnum cupressiforme var. filiforme may well be nothing more than an environmental modification but is retained as a variety until its position is clarified.
492
Hypnum subjulaceum was published by Schlesak et al. (2018) as Hypnum subcomplanatum rather than Hypnum subjulaceum because the latter was
thought to be blocked by the name Hypnum subjulaceum Besch. However, the latter name was not validly published, so its use at species level is not
blocked. This was corrected by Schlesak et al. (2019). Later, Kučera, Kuznetsova, et al. (2019) found that Hypnum subjulaceum formed a supported
lineage within Hypnum cupressiforme s.lat., and so it was returned to varietal status.
493
Hypnum resupinatum is treated at species level, as it is usually distinct morphologically and has a distinct geographical distribution. However, according to
current knowledge, it might equally well be treated as a variety of Hypnum cupressiforme, as in Guerra (2018). Ongoing molecular studies should clarify the
situation.
494
The new family Callicladiaceae was established by Kučera, Kuznetsova, et al. (2019) on molecular grounds.
495
Molecular evidence places Hypnum imponens in Callicladium (Schlesak et al. 2018).
496
The family Taxiphyllaceae was established by Ignatov, Afonina, et al. (2012) to accommodate Taxiphyllum.
497
Frahm (2013) synonymised Brotherella lorentziana with Brotherella henonii (Duby) M.Fleisch., an east Asian species, but a genetic review is still pending. We
therefore continue to accept Brotherella lorentziana as valid.
498
The name Hypnum affine Hook., basionym of Heterophyllium affine, is considered an orthographic variant of the older Hypnum adfine Hoffm. ex Schwägr.
(Allen 2018). The next available synonym is Hypnum nemorosum.
499
The new family Jocheniaceae was established by Kučera, Kuznetsova, et al. (2019) on molecular evidence.
500
Schlesak et al. (2018) established the genus Jochenia for Hypnum pallescens.
501
Using molecular data, Kučera, Kuznetsova, et al. (2019) found that Jochenia protuberans was consistently different from Jochenia pallescens. Although this taxon
has not often been recognised even at varietal level, there is now sufficient evidence to treat it as a species. Ando (1973) correctly distinguished between the taxa,
although he treated Jochenia protuberans only as a form of Jochenia pallescens. His synonymy of Hypnum reptile with Hypnum pallescens and clarification of the
misunderstanding concerning the type of Leskea pallescens Hedw. introduced by the authors of Bryologia Europaea, is followed here.
502
The family Stereodontaceae was established by Schlesak et al. (2018) to legitimate the high statistical support for the clade including Stereodon.
503
The treatment of Stereodon follows Schlesak et al. (2018) and Kučera, Kuznetsova, et al. (2019).
490
491
JOURNAL OF BRYOLOGY
1
2
3
4
5
6
S. aemulans (Breidl.) Broth. [Hypnum aemulans Breidl.]504
S. callichrous (Brid.) Lindb. [Hypnum callichroum Brid.]
S. hamulosus (Schimp.) Lindb. [Hypnum
hamulosum Schimp.]
S. holmenii (Ando) Ignatov & Igantova
[Hypnum holmenii Ando]
S. pratensis (W.D.J.Koch ex Spruce) Warnst.
[Breidleria pratensis (W.D.J.Koch ex
Spruce) Loeske]
S. subimponens (Lesq.) Broth. [Hypnum subimponens Lesq.]
1 P. procerrimus (Molendo) M.Fleisch. [Ctenidium procerrimum (Molendo) Lindb.,
Hypnum procerrimum Molendo]
272
Ptilium De Not.511
1 P. crista-castrensis (Hedw.) De Not.
273
Pylaisia Schimp.
1 P. polyantha (Hedw.) Schimp.
2 P. selwynii Kindb.
274
Roaldia P.E.A.S.Câmara & Carv.-Silva512
1 R. dolomitica (Milde) Hedenäs, Schlesak &
D.Quandt [Hypnum revolutum (Mitt.)
Lindb. var. dolomiticum (Milde) Mönk.]513
2 R. revoluta (Mitt.) P.E.A.S.Câmara & M.Carvalho-Silva [Hypnum revolutum (Mitt.)
Lindb.]
275
Vesicularia (Müll.Hal.) Müll.Hal.514
1 V. reimersiana Bizot & P.de la Varde
Pylaisiaceae Schimp.
266
267
Aquilonium Hedenäs, Schlesak & D.Quandt505
1 A. plicatulum (Lindb.) Hedenäs, Schlesak &
D.Quandt
[Hypnum
plicatulum
(Lindb.) A.Jaeger]
Buckia D.Rios, M.T.Gallego & J.Guerra506
1 B. vaucheri (Lesq.) D.Rios, M.T.Gallego &
J.Guerra [Hypnum vaucheri Lesq.]
268
Calliergonella Loeske
1 C. cuspidata (Hedw.) Loeske
2 C. lindbergii (Mitt.) Hedenäs
269
Homomallium (Schimp.) Loeske
1 H. incurvatum (Schrad. ex Brid.) Loeske
270
Pseudohygrohypnum Kanda507
1 P. eugyrium (Schimp.) Kanda [Hygrohypnum
eugyrium (Schimp.) Broth.]
2 P. fertile (Sendtn.) Jan Kučera & Ignatov
[Hypnum fertile Sendtn., Stereodon
fertilis (Sendtn.) Lindb.]508
3 P. subeugyrium (Renauld & Cardot) Ignatov &
Ignatova [Hygrohypnum subeugyrium
(Renauld & Cardot) Broth.]509
271
504
Pseudostereodon (Broth.) M.Fleisch.510
63
Sematophyllaceae Broth.
276 Sematophyllum Mitt.
1 S. adnatum (Michx.) E.Britton
2 S. demissum (Wilson) Mitt.
3 S. substrumulosum (Hampe) E.Britton
Hylocomiaceae M.Fleisch.
277 Hageniella Broth.515
1 H. micans (Mitt.) B.C.Tan & Y.Jia
278
Hylocomiadelphus Ochyra & Stebel516
1 H. triquetrus (Hedw.) Ochyra & Stebel [Rhytidiadelphus triquetrus (Hedw.) Warnst.]
279
Hylocomiastrum Broth.
1 H. pyrenaicum (Spruce) M.Fleisch.
2 H. umbratum (Hedw.) M.Fleisch.
280
Hylocomium Bruch & Schimp.
1 H. splendens (Hedw.) Schimp.
Hypnum aemulans has been treated as a synonym of Stereodon hamulosus (Hypnum hamulosum) by most European authors, and was therefore omitted by
Hill et al. (2006). No useful DNA could be extracted from this species during the recent revision of European Hypnum by Schlesak et al. (2018), so its generic
placement remains uncertain. However, morphological evidence suggests that it should be located in Stereodon.
505
The genus Aquilonium was established by Schlesak et al. (2018) to accommodate, among other species, Hypnum plicatulum.
506
The genus Buckia was established by Câmara et al. (2018) to accommodate Hypnum vaucheri.
507
Pseudohygrohypnum, described by Kanda (1976 (1977)), is accepted, as a broad concept of Hygrohypnum can no longer be maintained.
508
Kučera, Kuznetsova, et al. (2019) placed Hypnum fertile in Pseudohygrohypnum on molecular evidence.
509
Hygrohypnum subeugyrium was transferred to Pseudohygrohypnum by Ignatov and Ignatova (2004).
510
Pseudostereodon was re-established by Câmara et al. (2018).
511
The placement of Ptilium in the Pylaisiaceae is a temporary soluation, as its position remains uncertain until further research takes place.
512
The genus Roaldia was established to accommodate Hypnum revolutum s.lat. (Câmara et al. 2018).
513
While Roaldia dolomitica is regarded as ’mere morphological variation’ by Câmara et al. (2018), it is treated as a full species by Schlesak et al. (2018). The
latter view is supported by Kučera, Kuznetsova, et al. (2019).
514
The placement of Vesicularia in Pylaisiaceae follows Ignatov, Gardiner, et al. (2007).
515
Although placed in Hylocomiaceae by Frey and Stech (2009), Hageniella is probably best placed in Sematophyllaceae.
516
Hylocomiadelphus was pubished by Ochyra and Stebel (2008) to accommodate Rhytidiadelphus triquetrus. This was not widely accepted, but recent molecular work (Ignatov, Ignatova, et al. 2019) has supported the idea.
64
N. G. HODGETTS ET AL.
281
Loeskeobryum Broth.
1 L. brevirostre (Brid.) M.Fleisch.
282
Pleurozium Mitt.
1 P. schreberi (Willd. ex Brid.) Mitt.
283
Rhytidiadelphus (Limpr.) Warnst.
1 R. loreus (Hedw.) Warnst.
2 R. squarrosus (Hedw.) Warnst.
3 R. subpinnatus (Lindb.) T.J.Kop.
Rhytidiaceae Broth.
284 Rhytidium (Sull.) Kindb.
1 R. rugosum (Hedw.) Kindb.
Entodontaceae Kindb.
285 Entodon Müll.Hal.
1 E. challengeri (Paris) Cardot
2 E. cladorrhizans (Hedw.) Müll.Hal.
3 E. concinnus (De Not.) Paris
4 E. schleicheri (Schimp.) Demet.
Cryphaeaceae Schimp.
286 Cryphaea D.Mohr
1 C. heteromalla (Hedw.) D.Mohr
287
Dendrocryphaea Broth.
1 D. lamyana (Mont.) P.Rao
Leucodontaceae Schimp.
288 Leucodon Schwägr.
1 L. canariensis (Brid.) Schwägr.
2 L. flagellaris Lindb. ex Broth.517
3 L. immersus Lindb.518
4 L. pendulus Lindb.
5 L. sciuroides (Hedw.) Schwägr. [Leucodon
sciuroides var. morensis (Schwägr.)
De Not.]519
6 L. treleasei (Cardot) Paris
289
Nogopterium Crosby & W.R.Buck [Pterogonium
Sw. nom. illeg.]520
1 N. gracile (Hedw.) Crosby & W.R.Buck [Pterogonium gracile (Hedw.) Sm.]
Antitrichiaceae Ignatov & Ignatova
290 Antitrichia Brid.
517
1 A. californica Sull.
2 A. curtipendula (Hedw.) Brid.
Neckeraceae Schimp.
291 Alleniella S.Olsson, Enroth & D.Quandt521
1 A. besseri (Lobarz.) S.Olsson, Enroth &
D.Quandt [Neckera besseri (Lobarz.)
Jur.]
2 A. complanata (Hedw.) S.Olsson, Enroth &
D.Quandt
[Neckera
complanata
(Hedw.) Huebener]
292
Exsertotheca S.Olsson, Enroth & D.Quandt522
1 E. baetica (J.Guerra) Draper, GonzálezMancebo, O.Werner, J.Patiño & Ros
[Neckera baetica J.Guerra ]523
2 E. crispa (Hedw.) S.Olsson, Enroth & D.Quandt
[Neckera crispa Hedw.]
3 E. intermedia (Brid.) S.Olsson, Enroth &
D.Quandt [Neckera intermedia Brid.]
293
Homalia (Brid.) Bruch & Schimp.
1 H. lusitanica Schimp.
2 H. trichomanoides (Hedw.) Brid.
294
Iwatsukiella W.R.Buck & H.A.Crum524
1 I. leucotricha (Mitt.) W.R.Buck & H.A.Crum
295
Leptodon D.Mohr
1 L. corsicus Enroth, Sotiaux, D.Quandt &
Vanderp.525
2 L. longisetus Mont. [Cryptoleptodon longisetus
(Mont.) Enroth]526
3 L. smithii (Hedw.) F.Weber & D.Mohr
296
Neckera Hedw.
1 N. cephalonica Jur. & Unger
2 N. menziesii Drumm.
3 N. oligocarpa Bruch
4 N. pennata Hedw.
5 N. pumila Hedw.
297
Pseudanomodon (Limpr.) Ignatov & Fedosov527
1 P. attenuatus (Hedw.) Ignatov & Fedosov
[Anomodon
attenuatus
(Hedw.)
Huebener]
Werner, Rodriguez-Atienza, et al. (2015) confirmed the status of Leucodon flagellaris.
Werner, Rodriguez-Atienza, et al. (2015) confirmed the status of Leucodon immersus.
Werner, Rodriguez-Atienza, et al. (2015) found no substantial basis for retaining Leucodon sciuroides var. morensis.
520
Although familiar through long use, Pterogonium is an illegitimate name. It was replaced by Nogopterium (an anagram) by Crosby and Buck (2011).
521
Alleniella was segregated from Neckera by Olsson et al. (2011).
522
Exsertotheca was segregated from Neckera by Draper, González-Mancebo, et al. (2011).
523
Neckera baetica was described by Guerra, Jiménez-Martínez, Jiménez (2010) and transferred to Exsertotheca by Olsson et al. (2011).
524
Iwatsukiella has been placed in the Heterocladiaceae Ignatov & Ignatova, but this family name is illegitimate, having previously been used for a family of
algae. Therefore the genus is temporarily placed in Neckeraceae until another name is substituted.
525
Leptodon corsicus, apparently a Corsican endemic, was described by Sotiaux et al. (2009).
526
Cryptoleptodon longisetus was transferred to Leptodon by Olsson et al. (2011).
527
Pseudanomodon was established for Anomodon attenuatus after molecular studies showed it to be deeply nested in the Neckeraceae (Ignatov, Fedorova,
et al. 2019).
518
519
JOURNAL OF BRYOLOGY
298
Thamnobryum Nieuwl.
1 T. alopecurum (Hedw.) Gangulee
2 T. angustifolium (Holt) Nieuwl.528
3 T. cataractarum N.G.Hodgetts & Blockeel529
4 T. fernandesii Sérgio530
5 T. maderense (Kindb.) Hedenäs
6 T. neckeroides (Hook.) E.Lawton
7 T. rudolphianum Mastracci
8 T. subserratum (Hook. ex Harv.) Nog. &
Z.Iwats.531
Heterocladiellaceae Ignatov & Fedosov532
299 Heterocladiella Ignatov & Fedosov
1 H. dimorpha (Brid.) Ignatov & Fedosov [Heterocladium
dimorphum
(Brid.)
Schimp.]
5 I. montanum Draper, Hedenäs, M.Stech,
T.Lopes & Sim-Sim535
6 I. myosuroides Brid. [Isothecium myosuroides
subsp. brevinerve Kindb.]536
7 I. prolixum (Mitt.) M.Stech, Sim-Sim, Tangney
& D.Quandt [Echinodium prolixum
(Mitt.) Broth.]537
Echinodiaceae Broth.
302 Echinodium Jur.
1 E. renauldii (Cardot) Broth.
2 E. setigerum (Mitt.) Jur.
3 E. spinosum (Mitt.) Jur.
303
Pseudomalia Enroth538
1 P. webbiana (Mont.) Enroth [Homalia webbiana (Mont.) Schimp., Neckera webbiana (Mont.) Düll]
Lembophyllaceae Broth.
300 Heterocladium Bruch & Schimp.533
1 H. flaccidum (Schimp.) A.J.E.Sm.
2 H. heteropterum (Brid.) Schimp.
3 H. wulfsbergii I.Hagen
Myuriaceae M.Fleisch.539
304 Andoa Ochyra
1 A. berthelotiana (Mont.) Ochyra
301
305
Ctenidium (Schimp.) Mitt.
1 C. molluscum (Hedw.) Mitt.540
306
Hyocomium Bruch & Schimp.
1 H. armoricum (Brid.) Wijk & Margad.
307
Myurium Schimp.
1 M. hochstetteri (Schimp.) Kindb.
528
Isothecium Brid.
1 I. algarvicum W.E.Nicholson & Dixon
2 I. alopecuroides (Lam. ex Dubois) Isov.
3 I. holtii Kindb.
4 I. interludens Stirt. [Isothecium myosuroides
var.
brachythecioides
(Dixon)
534
Braithw.]
65
Furness and Gilbert (1980) showed that Thamnobryum angustifolium maintains its characters distinct from Thamnobryum alopecurum in culture. There are
at least three distinct morphological features that distinguish this species from Thamnobryum alopecurum. Hodgetts and Blockeel (1992) considered it to be
more closely related to Thamnobryum cataractarum and the Madeiran Thamnobryum fernandesii than to Thamnobryum alopecurum. However, more recent
molecular work by Olsson et al. (2009) suggests that, while Thamnobryum angustifolium is undoubtedly a distinct entity morphologically, colonies of this
plant (and of the other narrowly endemic Thamnobryum species) may originate from the surrounding subpopulations of Thamnobryum alopecurum. In this
case, the two subpopulations of Thamnobryum angustifolium are independently derived from local Thamnobryum alopecurum, and have evolved convergently in response to the rheophilous habitat (Blockeel et al. 2014).
529
Olsson et al. (2009) used molecular techniques to suggest that, like Thamnobryum angustifolium, Thamnobryum cataractarum may be a local derivative of
Thamnobryum alopecurum, evolving convergently in response to its habitat (Blockeel et al. 2014).
530
Recent molecular work by Olsson et al. (2009) suggests that while Thamnobryum fernandesii is a distinct entity, colonies may, like Thamnobryum angustifolium and Thamnobryum cataractarum, originate from surrounding colonies of Thamnobryum alopecurum.
531
Köckinger, Suanjak, et al. (2008), when reporting Thamnobryum subserratum from Austria, pointed out that it was first given for Europe by Mastracci
(2003), in which Figure 2 (l, m) was drawn from a collection from Latvia (Kurland, leg. Malta, Z.).
532
The new family Heterocladiellaceae was proposed to accommodate Heterocladium dimorphum. The taxonomic position of this species is still uncertain, but
the genetic differentiation was considered sufficient to place it in a separate genus and family close to Echinodiaceae, Lembophyllaceae and Neckeraceae
(Ignatov, Fedorova, et al. 2019).
533
Heterocladium has been placed in the Heterocladiaceae Ignatov & Ignatova, but this family name is illegitimate, having previously been used for a family of
algae. It was temporarily placed in the Neckeraceae until molecular evidence showed that it is best placed in the Lembophyllaceae (Ignatov, Fedorova, et al.
2019).
534
Hodgetts and Vanderpoorten (2018) returned Isothecium myosuroides var. brachythecioides to species level on the basis of molecular and morphological
evidence.
535
Isothecium montanum, a Madeiran endemic, was described by Draper, Hedenäs, et al. (2015).
536
The features characterising Isothecium myosuroides subsp. brevinerve are very plastic and have no taxonomic value. The authorship of this subspecies is
‘Kindb.’ rather than ‘Lindb.’, as it was listed in Hill et al. (2006).
537
Echinodium prolixum was transferred to Isothecium by Stech et al. (2008).
538
On the basis of molecular studies, Pseudomalia was established as a new genus in the Echinodiaceae to accommodate Homalia webbiana (Enroth et al.
2019), a species long suspected not to belong in the Neckeraceae (Olsson et al. 2009).
539
Andoa, Ctenidium and Myurium form a well supported clade in Huttunen, Bell, et al. (2012). Furthermore, these three genera plus Hyocomium form a clade
in Cox et al. (2010).
540
There continues to be no firm basis for any of the infraspecific taxa of Ctenidium molluscum, so the treatment of Hill et al. (2006) is continued.
66
N. G. HODGETTS ET AL.
Anomodontaceae Kindb.
308 Anomodon Hook. & Taylor [Anomodontella
Ignatov & Fedosov, Anomodontopsis
Ignatov & Fedosov, [Haplohymenium
Schwägr.]541
1 A. longifolius (Schleich. ex Brid.) Hartm.
[Anomodontella longifolia (Schleich.
ex Brid.) Ignatov & Fedosov]
2 A. rugelii (Müll.Hal.) Keissl. [Anomodontopsis
rugelii (Müll.Hal.) Ignatov & Fedosov]
3 A. tristis (Ces.) Sull. & Lesq. [Haplohymenium
triste (Ces.) Kindb.]
4 A. viticulosus (Hedw.) Hook. & Taylor
Taxa rejected from the covered area
The following taxa have been reported from the area
but have been shown not to occur there. Taxa reported
from ‘Europe’ without any further information and
which are clearly errors are not included in the list
below. There are about 150 liverwort names (mostly
subspecific names) with types from Europe that have
not been recognised recently, nor synonymized.
Some of those names are so old that they may threaten
accepted names if their true identity becomes known.
Some of the names in Jungermannia were enumerated
in Söderström, Hagborg, et al. (2016).
Liverworts
Acrobolbus anisodontus (Hook.f. & Taylor) Briscoe [Tylimanthus anisodontus (Hook.f. & Taylor) Mitt.] was
reported from Madeira by Stephani (1905) but rejected
as Acrobolbus madeirensis by Grolle and Persson (1966),
and from the Azores by Allorge and Allorge (1948) but
rejected as Acrobolbus azoricus.
Acrobolbos ciliatus (Mitt.) Schiffn. is reported from
Britain (coll. D. G. Long 29767, E) by Masuzaki et al.
(2010) but this is an error for Acrobolbus wilsonii
(D. G. Long, pers. comm.).
Acrobolbus laxus (Lehm. & Lindenb.) Briscoe is
reported from the Azores and Madeira by Burghardt
and Gradstein (2008) but their concept included Acrobolbus azoricus and Acrobolbus madeirensis to which
the reports belong.
Asterella blumeana (Nees) Kachroo is a southeast
Asian taxon reported as Fimbraria blumeana Nees
from Italy by Tassi (1901) but rejected by Aleffi and
Schumacker (1995).
Asterella elegans (Spreng.) Trevis. is a South American taxon reported from Corsica as Fimbraria elegans
541
Spreng by Casares-Gil (1919) but this is rejected by Bischler and Jovet-Ast (1973).
Asterella marginata (Nees) S.W.Arnell was reported
from Madeira as Fimbraria marginata Nees by Gottsche
et al. (1846) but this must be rejected. It is a southern
African taxon.
Asterella pringlei Underw. is an American taxon
reported from Italy as Fimbraria stahlii Steph. by Tassi
(1901) but this is rejected by Aleffi and Schumacker
(1995).
Asterella tenella (L.) P.Beauv. is a North American
taxon reported as Fimbraria tenella L. from several
places in Europe during the 19th centuary, but all
reports are rejected by Long (2006).
Asterella wallichiana (Lehm. & Lindenb.) Grolle is an E
Asian taxon reported from Italy as Fimbraria raddii
Corda ex Nees (type). However, the type is probably
from Himalaya and certainly not from Europe (Long
2006).
Bazzania denudata (Lindenb. & Gottsche) Trevis. is a
taxon from North America and East Asia reported many
times from central and western Europe but rejected
from here by Schuster (1969).
Blepharostoma arachnoideum M.Howe is a North
American taxon reported from Polar Ural by Zinovjeva
(1973) but rejected as B. trichophyllum subsp. brevirete
by Konstantinova and Potemkin (1994).
Cephalozia lucens (A.Evans) Steph. is a taxon from
Hawaii reported from Italy by Rhodegher (1896) but
rejected by Aleffi (2005).
Cephaloziella mammillifera R.M.Schust. & Damsh. is
an American taxon reported from ‘Europe’ by Stotler
and Crandall-Stotler (2017). This taxon was treated as
a questioned synonym of Cephaloziella varians var.
scabra by Damsholt (2002) and if this synonym is
correct, the taxon does occur in Europe.
Cephaloziella verrucosa Steph. is a taxon from the
Southern Hemisphere that has erroneously been
reported from Svalbard (Bryhn 1909), probably due to
nomenclatural confusion.
Cyathodium spurium (Dicks.) Lindb. ex Braithw. may
be conspecific with Cyathodium cavernarum
(Braithwaite 1878). The type specimen is from Scotland,
but it is probably mislabelled as no Cyathodium is
known from boreal Europe.
Frullania davurica Hampe ex Gottsche, Lindenb. &
Nees. has been reported several times from Europe,
mostly because it has sometimes included Frullania
jackii as a subspecies [F. davurica subsp. jackii
(Gottsche) S.Hatt.].
Frullania inflata Gottsche is shown to be a complex
species confined in a restricted sense to North
America while the European populations must be
Ignatov, Fedorova, et al. (2019) established the genera Anomodontopsis (for Anomodon rugelii) and Anomodontella (for Anomodon longifolius), and resurrected Haplohymenium (for Anomodon tristis). Although Ignatov, Fedorova, et al. (2019) show different lineages ‘within the Anomodontaceae’, the same
study gives support for a clade that includes all the species of Anomodon s.lat. except Anomodon attenuatus, so a relatively conservative approach is
adopted here in order to avoid the unnecessary proliferation of small genera.
JOURNAL OF BRYOLOGY
named Frullania cleistostoma (Mamontov, Potemkin,
et al. 2018).
Frullania muscicola Steph. is an Asiatic taxon
reported from Europe as a putative synonym (variety)
of F. cesatiana (=Frullania inflata; see note under the
latter).
Frullania obscurifolia Mitt. is an African taxon
reported by Dirkse et al. (1993) but rejected as Frullania
azorica by González-Mancebo et al. (2008).
Herbertus dicranus (Gottsche, Lindenb. & Nees)
Trevis. has (sometimes as Herbertus sakuraii (Warnst.)
S.Hatt.) been reported from Norway and Scotland (Konstantinova 2000a; Hodgetts 2003) due to the inclusion
of the now rejected synonymy with Herbertus borealis.
The unsupported mention of ‘Madeira’ (Juslén 2006)
is an obvious error for Madagascar.
Heteroscyphus integrifolius (Lehm. & Lindenb.)
Fulford is a taxon from South America reported from
Italy by Rodegher (1896) but rejected by Aleffi (2005).
Jubula hutchinsiae subsp. javanica (Steph.) Verd. is
an Asiatic taxon. It was supposed to be the subspecies
occurring in Caucasus prior to the description of subsp.
caucasica. It has also erroneously been reported from
other parts of Europe and Macaronesia.
Jubula hutchinsiae subsp. pennsylvanica (Steph.)
Verd. is the North American subspecies that has been
reported erroneously from Ireland, France, Macaronesia
and Caucasus.
Jungermannia exsertifolia Steph. is an East Asiatic
species reported from Europe on the basis that it
includes Jungermannia eucordifolia as a subspecies,
but Mamontov, Konstantinova, et al. (2018) showed
that the latter deserves specific rank.
Kurzia makinoana (Steph.) Grolle is a species from
North America and East Asia that has been reported
from several places in Europe since Grolle (1963)
included Kurzia sylvatica as synonym. However, this
synonymy was rejected by Grolle (1973).
Lejeunea laetevirens Nees & Mont. is a primarly South
American taxon but with close affinity to Lejeunea
canariensis. Unless conspecific with the latter, Lejeunea
laetevirens is not found in Macaronesia.
Lophocolea humistrata (Hook.f. & Taylor) Gottsche,
Lindenb. & Nees is endemic to St Helena but was
noted also from Madeira by Stephani (1907).
Lophoziopsis propagulifera (Gottsche) Konstant. &
Vilnet is a Southern Hemisphere taxon that was considered conspecific with Lophoziopsis latifolia by
Bakalin (2005) but the synonymy was rejected by Köckinger (2017) and this is supported by ongoing unpublished molecular research by N. Konstantinova et al.
Metzgeria temperata Kuwah. is a primarily E Asiatic
taxon that has been recorded from all around the
Northern Hemisphere, including Europe and Macaronesia. However, the European and E Asiatic specimens
are not identical (cf. Köckinger 2017) and for the
67
present the name Metzgeria consanguinea is used
(see note also under that name).
Microlejeunea diversiloba (Spruce) Müll.Frib. is an
American taxon that was described from Mexico
under the name Lejeunea diversifolia Gottsche and
reported from Ireland by Spruce (1876a). However,
there exists an older Lejeunea diversifolia Mitt. (from
the Himalaya), and Spruce (1876b) changed the name
to Lejeunea diversiloba. It was subsequently reported
from the Azores and Ireland by Allorge and Persson
(1938). Grolle (1975) concluded that the Irish material
was not the same as the Mexican type and re-described
it as Lejeunea hibernica. All reports outside America
should belong here.
Plagiochila javanica (Sw.) Nees & Mont. is a taxon
from southeast Asia that Gottsche et al. (1844) reported
from the Canary Islands (leg. Webb) but this was
rejected by Inoue (1969) who could not locate any
specimen supporting the report.
Plagiochila patula (Sw.) Lindenb. is an American
species reported from Canary Islands (Dirkse et al.
1993) and Madeira (Nieuwkoop and Arts 1995), both
as Plagiochila dubia Lindenb. & Gottsche, but this was
rejected as Plagiochila virginica by Heinrichs, Pröschold
et al. (2002).
Plagiochila uniformis Mitt. is a taxon from southeast
Asia reported by Stephani (1903) as Plagiochila ambagiosa Mitt. from Ireland. Stephani’s concept of
P. ambagiosa was, however, not the same as the
taxon described by Mitten, but rather belonging to Plagiochila spinulosa.
Porella navicularis (Lehm. & Lindenb.) Pfeiff. is an
American taxon reported from several places in
central Europe, mostly during the 19th century, but
also a few times more recently.
Porella platyphylloidea (Schwein.) Lindb. is frequently reported from Europe. It has sometimes been
considered synonymous with Porella platyphylla but
the study by Heinrichs, Kreier, et al. (2011) shows that
the European and American populations of Porella platyphylla/platyphylloidea are different and only Porella
platyphylla occurs in Europe.
Riccia bullosa Link is described with two syntypes,
one from South Africa and the other from Portugal.
The lectotype is from South Africa and it is not conspecific with the European taxon (Grolle and Long
2000), the latter being Exormotheca welwitschii. All
reports from Europe are based on this confusion.
Riccia concava Bisch. ex C.Krauss is a taxon from
southern Africa reported from the Canary Islands as
Riccia capensis Steph. by Arnell (1961) and Eggers
(1982) but rejected by Perold (1989a).
Riccia limbata Bisch. ex C.Krauss is another
taxon from southern Africa reported from the Canary
Islands by Arnell (1961) and Eggers (1982) but this is
rejected by Perold (1989b) as Riccia nigrella.
68
N. G. HODGETTS ET AL.
Riccia oerstediana Lindenb. & Hampe is an American
taxon that Schuster (1992b, as Riccia stenophylla
Spruce) assumed was the taxon reported as fertile
Riccia fluitans from Europe. However, this has never
been verified.
Riella parisii Gottsche is reported twice from Europe,
Andalucia in Spain (Müller 1953) and in SE France
(Skrzypczak 2001). The Spanish report was rejected
by Brugués et al. (2011) and the French report by
Hugonnot (2019). It is now known only from Algeria
and Tunisia.
Sphaerocarpos texanus Austin is reported as widely
distributed in Europe but it is shown by Bell et al.
(2013) that the European populations are genetically
very distinct from the American populatiopns, although
the morphological differences are small. Sphaerocarpos
europaeus should be used for the European species.
Telaranea nematodes (Gottsche ex Austin) M.Howe is
a neotropical-tropical African species which in the past
has been reported from several places in southwestern Europe and Macaronesia but Engel and
Smith Merrill (2004) showed that the European and
Macaronesian populations represent a different taxon,
Telaranea europaea.
Telaranea sejuncta (Ångstr.) S.W.Arnell is an American
species that has been treated as a synonym of Telaranea
nematodes and reported from Europe under this name.
Those reports represents Telaranea europaea.
Mosses
Amphidium tortuosum (Hornsch.) Cufod. is a widespread species in the tropics. The Macaronesian
endemic Amphidium curvipes was synonymised with
Amphidium tortuosum by Frahm et al. (2000), but SimSim et al. (2017) showed that Amphidium curvipes is
indeed distinct.
Barbula indica (Hook.) Spreng. is a widespread tropical species, which has been transferred to Hydrogonium, as H. orientale (F.Weber) Kučera, by Kučera,
Košnar, et al. (2013). All European records of ‘B. indica’
refer to H. consanguineum var. kurilense (Kučera,
Košnar, et al. 2013).
Bartramia stricta Brid. is considered by Damayanti
et al. (2012) to be restricted to South America. See footnote on Bartramia aprica above.
Brachymenium commutatum (Müll.Hal.) A.Jaeger is
excluded following a revision of the specimens from
Europe by Ros, Rams, et al. (2007).
Brachytheciastrum fendleri (Sull.) Ochyra & Żarnowiec
is a North American species; all European records of
B. fendleri refer to B. collinum (Orgaz et al. 2013).
Brotherella henonii (Duby) M.Fleisch. is an east Asian
species; molecular studies are needed to support the
proposed synonymy of Brotherella lorentziana (Frahm
2013) with this species.
Bryum philonotulum Müll.Hal. is a synonym of Pohlia
philonotula (Müll.Hal.) Broth., a little-known African
species, and not to be confused with Bryum philonotulum Hampe, listed here as a synonym of Bryum
kikuyense.
Cinclidotus pachylomoides Bizot is a mainly Asian
species; European records have not been confirmed
and it is considered very doubtful that this species
occurs in Europe.
Drepanocladus longifolius (Mitt.) Paris is an exclusively Southern Hemisphere species; Saługa et al.
(2018) showed that European records of this species
should be referred to Drepanocladus capillifolius.
Lindbergia brachyptera (Mitt.) Kindb. occurs only in
North America (Ignatova, Ignatov, et al. 2010).
Pohlia saprophila (Müll.Hal.) Broth. has been
removed from European list as the European Russian
records were based on misidentifications of Pohlia
longicolla (Czernyadjeva et al. 2017).
Sanionia georgicouncinata (Müll.Hal.) Ochyra &
Hedenäs is a Southern Hemisphere species distinct
from S. nivalis (Hedenäs 2012).
Sphagnum lescurii Sull. is a North American species
that has been reported from Europe in the past (e.g.
by Corley et al. 1981). European records refer to Sphagnum auriculatum and Sphagnum inundatum.
Sphagnum magellanicum Brid. has been shown to be
confined to southern South America (Hassel et al. 2018).
Tortella arctica (Arnell) Crundw. & Nyholm has not
been reliably reported from Europe; see note on T. x
cuspidatissima above.
Taxa only occurring in areas adjacent to the
covered area
The following taxa have not yet been found in Europe
or northern Macaronesia, but occur just outside the
area (in Cape Verde, North Africa, Turkey, Georgia, Azerbadjan, Armenia or western Siberia). Information for
liverworts has been obtained from the ELPT database
(Söderström, Hagborg, et al. 2019) and for mosses
mainly from Ignatov, Afonina, et al. (2006), Kürschner
and Erdağ (2005), O’Shea (2006 with updates) and
Ros, Mazimpaka, et al. (2013). Some of these taxa are
obscure and known only from the original collections;
others are mainly American or Pacific species which
need to be confirmed in the territories listed.
Liverworts
Acrolejeunea emergens (Mitt.) Steph. (Cape Verde)
Cheilolejeunea xanthocarpa (Lehm. & Lindenb.)
Malombe (Cape Verde)
Cyathodium cavernarum Kunze (Cape Verde)
Frullania socotrana Mitt. (Cape Verde)
Frullania spongiosa Steph. (Cape Verde)
Lejeunea capensis Gottsche (Cape Verde)
JOURNAL OF BRYOLOGY
Lophozia lacerata N.Kitag. (Georgia)
Lophoziopsis excisa var. infuscata (R.M.Schust. &
Damsh.) Konstant. & Vilnet (Yamal-Nenets)
Lophoziopsis excisa var. succulenta (R.M.Schust. &
Damsh.) Konstant. & Vilnet (Yamal-Nenets)
Marchantia debilis K.I.Goebel (Morocco, Egypt)
Marchantia pappeana Lehm. subsp. pappeana (Cape
Verde)
Plagiochasma eximium (Schiffn.) Steph. (Cape Verde)
Plagiochasma microcephalum (Steph.) Steph. var. tunesicum Bischl. (Tunisia)
Pleurozia gigantea (F.Weber) Lindb. (Cape Verde)
Prasanthus jamalicus Potemkin (Yamal-Nenets)
Radula prolifera Arnell (Yamal-Nenets)
Riccia chudoana Steph. (Algeria)
Riccia congoana Steph. (Egypt)
Riccia mamillata Trab. ex Steph. (Algeria)
Riccia polycarpa (Trab.) Jelenc (Algeria)
Riella cyrenaica Maire (Libya)
Riella numidica Trab. (Morocco, Algeria, Libya)
Riella parisii Gottsche (Algeria, Tunisia)
Riella sersuensis Trab. (Algeria)
Scapania brevicaulis Taylor (western Siberia)
Scapania cuspiduligera var. diplophyllopsis R.M.Schust.
(Yamal-Nenets)
Scapania microdonta (Mitt.) Müll.Frib. (Sverdlovsk)
Solenostoma lignicola (Schiffn.) Váňa, Hentschel & Heinrichs (Turkey)
Solenostoma subtilissimum (Schiffn.) R.M.Schust.
(Turkey)
Syzygiella manca (Mont.) Steph. (Cape Verde)
Mosses
Acaulon longifolium Herrnst. & Heyn (Israel)
Aulacomnium acuminatum (Lindb. & Arnell) Kindb.
(Arctic Western Siberia)
Brachymenium acuminatum Harv. (Cape Verde)
Brachymenium commutatum (Müll.Hal.) A.Jaeger
(Algeria, Tunisia)
Brachymenium exile (Dozy & Molk.) Bosch. & Sande Lac.
(Algeria, Cape Verde, Lebanon)
Brachytheciastrum bellicum (W.R.Buck, J.A.Jiménez, Ros
& M.J.Cano) Vanderp., Ignatov, Huttunen & Goffinet
(Morocco)
Brachytheciastrum umbilicatum (Jur. & Milde) Orgaz,
M.J.Cano & J.Guerra (Turkey)
Bryum anomodon Mont. (Cape Verde)
Bryum atrovirens Brid. (Turkey)
Calohypnum plumiforme (Wilson) Jan Kučera & Ignatov
(Georgia)
Cinclidotus bistratosus Kürschner & Lübenau-Nestle
(Turkey)
Cinclidotus pachyloma E.S.Salmon (Israel, Lebanon,
Turkey)
Cinclidotus vardaranus Erdağ & Kürschner (Turkey)
Crumia latifolia (Kindb.) W.B.Schofield (Armenia)
69
Didymodon caboverdeanus J.A.Jiménez & M.J.Cano
(Cape Verde)
Didymodon tectorum (Müll.Hal.) K.Saito (Egypt)
Entodon pseudoseductrix (Müll.Hal.) A.Jaeger (Cape
Verde)
Entodontopsis leucostega (Brid.) W.R.Buck & Ireland
(Cape Verde)
Entosthodon angustifolius Jur. & Milde (Jordan, Libya,
Turkey)
Entosthodon niloticus Schimp. (Egypt)
Epipterygium rigidum Lindb. ex Broth. (Georgia)
Erpodium grossirete Müll.Hal. (Cape Verde)
Erpodium perrottetii (Mont.) A.Jaeger & Sauerb. (Cape
Verde)
Fabronia gueinzii Hampe (Morocco)
Fabronia leikipiae Müll.Hal. (Cape Verde)
Fissidens allorgei P.de la Varde (Cape Verde)
Fissidens androgynus Bruch ex C.Krauss (Cape Verde)
Fissidens dankelmannii Müll.Hal. (Cape Verde)
Fissidens flaccidus Mitt. (Cape Verde)
Fissidens megalotis Schimp. ex Müll.Hal. subsp. helictocaulos (Müll.Hal.) Brugg.-Nann. (Cape Verde)
Fissidens sciophyllus Mitt.(Cape Verde)
Fissidens usambaricus Broth. (Cape Verde)
Fontinalis antipyretica var. heldreichii (Müll.Hal.) Ruthe
(Turkey)
Funaria altissima Dixon (Algeria)
Funaria chevalieri P.de la Varde (Cape Verde)
Groutiella tomentosa (Hornsch.) Wijk & Marg. (Cape
Verde)
Gymnostomiella erosula (Müll.Hal. ex Dusén) Arts (Cape
Verde)
Gymnostomiella vernicosa (Hook. ex Harv.) M.Fleisch.
(Cape Verde)
Herpetineuron toccoae (Sull. & Lesq.) Cardot (Cape
Verde)
Hookeria acutifolia Hook. & Grev. (Georgia, Turkey)
Hydrogonium arcuatum (Griff.) Wijk & Margad. (Egypt)
Hydrogonium orientale (F.Weber) Jan Kučera (Cape
Verde, Egypt)
Hymenostylium congoanum Dixon & Naveau (Cape
Verde)
Hymenostylium hildebrandtii (Müll.Hal.) R.H.Zander
(Morocco)
Leptodictyum kurdicum (Schiffn.) Broth. (Turkey)
Leucodon bowringii Mitt. (Turkey)
Leucodon coreensis Cardot (Turkey)
Molendoa seravschanica Broth. & Györffy (Arctic
Western Siberia)
Orthotrichum urnaceum Müll.Hal. (Armenia, Azerbaijan)
Palamocladium leskeoides (Hook.) E.Britton (Cape Verde)
Perssonia sanguinea Bizot (Cape Verde)
Philonotis laxitexta J.Fröhl. (Lebanon)
Philonotis nanothecioidea Paris & Broth. (Cape Verde)
Physcomitrium immersum Sull. (Egypt)
Physcomitrium niloticum (Delile) Müll.Hal. (Egypt)
Platygyrella densa (Hook.) W.R.Buck (Cape Verde)
70
N. G. HODGETTS ET AL.
Plaubelia sprengelii (Schwägr.) R.H.Zander (Egypt)
Pohlia alba Lindb. & Arnell (Western Siberia)
Pohlia saprophila (Müll.Hal.) Broth. (Kazakhstan)
Pseudoleskeopsis bollei (Broth. & Geh.) P.Rao (Cape
Verde)
Pseudoleskeopsis pseudoattenuata (Müll.Hal.) Thér.
(Cape Verde)
Rhizomnium striatulum (Mitt.) T.J.Kop. (Turkey)
Schistidium cinclidodonteum (Müll.Hal.) B.Bremer
(Morocco)
Sphagnum lescurii Sull. & Lesq. (Turkey, if correct; this is
a North American species)
Sphagnum perfoliatum L.I.Savicz (Western Siberia and
Arctic Western Siberia)
Splachnobryum aquaticum Müll.Hal. (Jordan)
Splachnobryum limbatum D.H.Norris & R.H.Zander
(Egypt)
Syntrichia amphidiacea (Müll.Hal.) R.H.Zander (Cape
Verde)
Syntrichia caninervis var. pseudodesertorum (Vondr.)
M.T.Gallego (Turkey)
Tortella malacophylla (Müll.Hal.) Paris (Turkey)
Tortula acaulon var. galilaea (Herrnst. & Heyn) Ros &
Herrnst. (Israel)
Tortula grandiretis Broth. (Turkey)
Tortula kneuckeri Broth. & Geh. (Egypt)
Tortula plinthobia (Sull. & Lesq.) Broth. (Egypt)
Weissia breutelii Müll.Hal. (Israel, Turkey)
Weissia leptocarpa Schimp. ex Besch. (Turkey, if
accepted as a species – see annotation 160 in Hill
et al. 2006)
Weissia ovatifolia Kürschner (Jordan)
Weissia sinaloensis E.B.Bartram (Egypt)
Acknowledgements
Thanks are due to the large number of bryologists whose
research, expertise and experience has been drawn on in
the preparation of this checklist.
This project would not have been possible, or at least
would have been much more difficult, without the initial
Red List project, which was financed chiefly through an EC
LIFE project. All the bryologists who attended the five workshop meetings or otherwise answered our questions or
helped in any way during that project have contributed
directly or indirectly to this checklist:
Elvira Baisheva, Ariel Bergamini, Irene Bisang, Patrizia
Campisi, Annalena Cogoni, Tomas Hallingbäck, Marko Sabovljević, Norbert Schnyder, Cecilia Sérgio, Olga Afonina, Rosalina
Gabriel, César Garcia, Juana González Mancebo, Irina Goldberg,
Anders Hagborg, Sanna Huttunen, Elena Ignatova, Marta
Infante, Riikka Juutinen, Thomas Kiebacher, Niklas Lönnell,
Michael Lüth, Anabela Martins, Oleg Maslovsky, Gordon
Rothero, Sorin Ştefa nuţ, Kimmo Syrjänen, Alain Untereiner,
̵
Jiri VáňaI, Alain Vanderpoorten, Kai Vellak, Michele Aleffi, Jeff
Bates, Maude Baudraz, Des Callaghan, Nils Cronberg, Jo
Denyer, Jeff Duckett, Heinjo During, Vladimir Fedosov, Anna
Ganeva, Piotr Gorski, Urban Gunnarsson, Kristian Hassel,
Helena Hespanhol, Mark Hill, Rory Hodd, Kristofer Hylander,
Nele Ingerpuu, Sanna Laaka-Lindberg, David Long, Vicente
Mazimpaka, Anna Mežaka, Frank Müller, Jose David Orgaz,
Jairo Patiño, Sharon Pilkington, Felisa Puche, Rosa M. Ros,
Fred Rumsey, J. G. Segarra-Moragues, Ana Séneca, Adam
Stebel, Risto Virtanen, Henrik Weibull, Jo Wilbraham and Jan
Żarnowiec.
We are grateful to the following current or former IUCN
staff, who led the Red List workshop meetings and did a lot
of behind-the-scenes work: David Allen, Marta Cálix, Eve Englefield, Catarina Ferreira, Nicholas Fettes, Mariana García
Criado and Ana Nieto.
NGH would like to give special thanks to the Irish National
Parks and Wildlife Service for providing funding for a year to
work on the checklist and to Ed Wymer of the Irish National
Parks and Wildlife Service for doing a useful computer check
that resulted in the elimination of several errors and inconsistencies. The NTNU, Trondheim, and National Parks and Wildlife
Service, Department of Culture, Heritage & the Gaeltacht,
Dublin are thanked for funding the Open Access. We are grateful to Phil Stanley for all his help with compiling the index.
ORCID
L. Söderström
http://orcid.org/0000-0002-9315-4978
N.E. Bell
http://orcid.org/0000-0002-2401-2968
R. Garilleti
http://orcid.org/0000-0002-5977-2908
L. Hedenäs
http://orcid.org/0000-0003-1763-1696
J. Kučera
http://orcid.org/0000-0002-0230-5997
F. Lara
http://orcid.org/0000-0002-1665-5277
References
Afonina OM, Ignatova EA, Fedosov VE, Kuznetsova, OI. 2014.
Toward a new understanding of Syntrichia submontana
(Pottiaceae, Bryophyta). Arctoa. 23:11–24.
Aigoin DA, Huttunen S, Ignatov MS, Dirkse GM, Vanderpoorten
A. 2009. Rhynchostegiella (Brachytheciaceae): molecular recircumscription of a convenient taxonomic repository.
Journal of Bryology. 31:213–221.
Aleffi M. 2005. New check-list of the Hepaticae and
Anthocerotae of Italy. Flora Mediterranea. 15:485–566.
Aleffi M, Schumacker R. 1995. Check-list and Red-List of the
liverworts
(Marchantiophyta)
and
hornworts
(Anthocerotophyta) of Italy. Flora Mediterranea. 5:73–161.
Allen BH. 2014. Maine mosses: DrummondiaceaePolytrichaceae. Memoirs of the New York Botanical
Garden. 111:1–607.
Allen BH. 2018. Moss Flora of Central America. Part 4.
Fabroniaceae–Polytrichaceae. Monographs in Systematic
Botany from the Missouri Botanical Garden. 132: i-x, 1–830.
Allorge V, Allorge P. 1948. Végétation bryologique de l’Île de
Flores (Açores). Revue Bryologique & Lichénologique.
17:126–164.
Allorge P, Persson H. 1938. Contribution à la flore
hépaticologique des Îles Açores. Annales Bryologici. 11:6–14.
Alonso M, Jiménez JA, Cano MJ. 2018. New synonyms and
typifications in Chionoloma tenuirostre (Pottiaceae,
Bryophyta). Phytotaxa. 373:147–154.
Alonso M, Jiménez JA, Cano MJ. 2019. Taxonomic revision of
Chionoloma (Pottiaceae, Bryophyta). Annals of the Missouri
Botanical Garden. 104:563–632.
Alonso M, Jiménez JA, Nylander S, Hedenäs L, Cano MJ. 2016.
Disentangling generic limits in Chionoloma, Oxystegus,
Pachyneuropsis and Pseudosymblepharis (Bryophyta:
Pottiaceae): an inquiry into their phylogenetic relationships. Taxon. 65:3–18.
JOURNAL OF BRYOLOGY
Ando H. 1973. Studies on the genus Hypnum Hedw. (II).
Journal of Science of the Hiroshima University, series B, division 2. 14:165–207.
Andrus RE. 2006. Six new species of Sphagnum (Bryophyta:
Sphagnaceae). Sida. 22:959–972.
Aranda SC, Gradstein SR, Patiño J, Laenen B, Désamoré A,
Vanderpoorten A. 2014. Phylogeny, classification and
species
delimitation
in
the
liverwort
genus
Odontoschisma (Cephaloziaceae). Taxon. 63:1008–1025.
Arnell SW. 1961. List of hepaticae of the Canary Islands.
Svensk Botanisk Tidskrift. 55:379–393.
Bączkiewicz A, Szczecińska M, Sawicki J, Stebel A, Buczkowska
K. 2017. DNA barcoding, ecology and geography of the
cryptic species of Aneura pinguis and their relationships
with Aneura maxima and Aneura mirabilis (Metzgeriales,
Marchantiophyta). PLOS one. 12: e0188837:1–21.
Bakalin VA. 2001. Notes on Lophozia III. Some taxonomic problems in Lophozia sect. Lophozia. Arctoa. 10:207–218.
Bakalin VA. 2005. Monograficheskaia obrabotka roda
Lophozia (Dumort.) Dumort. s. str. [Monograph of the
genus Lophozia (Dumort.) Dumort. s. str.]. Moscow:
Nauka. Russian.
Bakalin VA. 2016. Notes on Lophozia VIII. The Lectotypification
of Lophozia longiflora (Nees) Schiffn. (Lophoziaceae,
Hepaticae). Herzogia. 29:635–643.
Bakalin VA, Fedosov VE, Fedorova AV, Nguyen VS. 2019.
Integrative
taxonomic
revision
of
Marsupella
(Gymnomitriaceae, Hepaticae) reveals neglected diversity
in Pacific Asia. Cryptogamie, Bryologie 40:59–85.
Bakalin VA, Klimova KG 2016. A note on Nardia japonica
Steph. (Gymnomitriaceae). Botanica Pacifica. 5:43–50.
Bakalin VA, Vilnet AA. 2018. A review of the genus
Diplophyllum (Marchantiophyta) in North and East Asia
with the description of a new species (D. sibiricum) based
on integrative taxonomy. Plant Systematics and
Evolution. 304:1269–1287.
Bakalin VA, Vilnet AA. 2019. Lophozia fuscovirens sp. nov.
(Lophoziaceae, Marchantiophyta): the second taxon with
brown gemmae within Lophozia s.s. Nordic Journal of
Botany. 37:1–10.
Bell D, Long DG, Forrest AD, Hollingsworth ML, Blom HH,
Hollingsworth PM. 2012. DNA barcoding of European
Herbertus (Marchantiopsida, Herbertaceae) and the discovery and description of a new species. Molecular Ecology
Resources. 12:36–47.
Bell D, Long DG, Hollingsworth P. 2013. The use of DNA barcoding to address major taxonomic problems for rare British
bryophytes. Edinburgh: Royal Botanic Garden.
Bell NE, Hyvönen J. 2010. A phylogenetic circumscription of
Polytrichastrum (Polytrichaceae): reassement of sporophyte morphology supports molecular phylogeny.
American Journal of Botany. 97:566–578.
Biehler JFP. 1807. Plantarum Novarum ex Herbario Sprengelii
Centuriam. Halis Saxonum: F.A. Grunerti Patr.
Bijlsma R, Van der Velde M, Van de Zande L, Boerema AC, Van
Zanten BO. 2000. Molecular markers reveal cryptic species
within Polytrichum commune (common hair-cap moss).
Plant Biology. 2:408–414.
Bisang I, Schumacker R, Sérgio C, Grolle R. 1989. Clé d’identification des espèces du genre Frullania Raddi (Hepaticae) en
Europe & en Macaronésie. Nuovo Giornale Botanico
Italiano. 122:255–266.
Bischler H, Jovet-Ast S. 1973. Les Hépatiques de Corse.
Énumeration, notes écologiques & biogéographiques.
Revue Bryologique & Lichénologique. 39:43–153.
Blackstock TH, Blackhall-Miles R, Ruffino L, Watling M. 2019.
Tricholepidozia
(Telaranea)
lindenbergii
(Gottsche)
71
E.D.Cooper (Marchantiophyta: Lepidoziaceae), another
antipodean liverwort established in Britain. Journal of
Bryology. 41:278–280.
Blockeel TL. 2017. The moss year – 2016. Field Bryology.
117:87–103.
Blockeel TL, Bosanquet SDS, Hill MO, Preston CD, editors.
2014. Atlas of British and Irish bryophytes. Newbury:
Pisces Publications.
Blockeel TL, Kučera J, Fedosov VE. 2017. Bryoerythrophyllum
duellii Blockeel (Bryophyta: Pottiaceae), a new moss
species from Greece and Cyprus, and its molecular
affinities. Journal of Bryology. 39:247–254.
Blom HH, Bednarek-Ochyra H, Ochyra R. 2016. Studies on
Schistidium (Grimmiaceae, Bryophyta) in Europe, with particular reference to the Alps: I. A description of S. marginale
sp. nov. Phytotaxa. 247:210–218.
Blom HH, Ignatova EA, Afonina OM. 2006. New records of
Schistidium (Grimmiaceae, Musci) in Russia. Arctoa.
15:187–194.
Bonfim Santos M, Stech M. 2017. Testing hypotheses on
suprageneric relationships and morphological evolution
in the Leucobryaceae (Bryophyta). Plant Systematics and
Evolution. 303:1383–1397.
Borovichev EA. 2008. New liverwort records from Мurmansk
Province. 1. Arctoa 17:191.
Borovichev EA, Mamontov YS. 2017. Hepaticae Rossicae
Exsiccatae. Fasc. XIII. Apatity: Russian Academy of Science,
Kola Science Centre, Polar-Alpine Botanical Garden Institute.
Bosanquet SDS, Lara F. 2012. Orthotrichum cambrense sp. nov.
(Orthotrichaceae), a distinctive moss from Wales, United
Kingdom. Cryptogamie, Bryologie. 33:329–339.
Braithwaite R. 1878. Riccia spuria, Dickson. Journal of Botany,
British and Foreign. 16:55.
Bridel SE. 1827. Bryologia Universa, seu systematica ad novam
methodum dispositio, historia et descriptio omnium muscorum frondosorum huscusque cognitorum cum synonymia ex auctoribus probatissimis. Vol. 1, pars 2. Leipzig: J.
A. Barth.
Brugués M, Infante M, Cros RM. 2011. Anotaciones sobre
hepáticas de España. Boletín de la Sociedad Española de
Biología. 36:3–8.
Brugués M, Sérgio C. 2010. On the identity of some
Entosthodon species endemic to Macaronesia or the
north of Africa. Cryptogamie, Bryologie. 31:277–280.
Bryhn N. 1909. Bryophyta pro flora Spitsbergensi nova. Nyt
Magazin for Naturvidenskaberne. 47:207–208.
Buch H. 1929. Eine neue moossystematische Methodik. In:
Winge, Ø, editor. Beretning om, det 18. Skandinaviske
naturforskermøde i København. 26–31 August 1929.
København: Frederiksberg Bogtrykkeri; p. 225–229.
Buch H. 1932. Vorarbeiten zu einer Lebermoosflora
Fenno-scandias. I. Ein Versuch zur Aufspaltung der
Gattungen Lophozia Dum. und Sphenolobus Steph.
Memoranda Societatis pro Fauna & Flora Fennica.
8:282–297.
Buchbender V, Hespanhol H, Krug M, Sérgio C, Séneca A, Mau
K, Hedenäs L, Quandt D. 2014. Phylogenetic reconstructions of the Hedwigiaceae reveal cryptic speciation and
hybridisation in Hedwigia. Bryophyte Diversity and
Evolution. 36:1–21.
Buczkowska K, Bakalin VA, Bączkiewicz A, Aguero B, Gonera P,
Ślipiko M, Szczecińska M, Sawicki J. 2018. Does Calypogeia
azurea (Calypogeiaceae, Marchantiophyta) occur outside
Europe? Molecular and morphological evidence. PLoS
One. 13:1–27.
Burghardt M, Gradstein SR. 2008. A revision of Tylimanthus
(Acrobolbaceae, Marchantiophyta) in Tropical America,
72
N. G. HODGETTS ET AL.
Africa, and Macaronesia. Fieldiana: Botany (n.ser.). 47:199–
210.
Burrell WH. 1911. Lophozia schultzii (Nees) Schiffn.) var. nov.
laxa. Journal of Botany. 49:217–219.
Callaghan DA. 2019. Typification and diagnosis of Weissia x
mittenii (Bruch & Schimp.) Mitt. emend. A.J.E.Sm. (Weissia
multicapsularis x W rostellata) (Pottiaceae, Bryophyta).
Journal of Bryology. 41:243–248.
Callaghan DA, Bell NE, Forrest LL. 2019. Taxonomic notes on
Weissia subgenus Astomum, including Weissia wilsonii
D.A.Callaghan, a new species from Europe. Journal of
Bryology. 41:135–148.
Callaghan DA, Masson J, During H. 2020. Physcomitrium × stevensoni D.A.Callaghan (Physcomitrium patens × P. eurystomum) (Funariaceae, Bryophyta), a new name for a rarely
recorded hybrid moss. Journal of Bryology. 42. https://
doi.org/10.1080/03736687.2020.1725325
Câmara PEAS, Carvalho-Silva M, Henriques DK, Guerra J,
Gallego MT, Poveda DR, Stech M. 2018. Pylaisiaceae
Schimp. (Bryophyta) revisited. Journal of Bryology.
40:251–264.
Cano MJ. 2007. Typification of the names of some infraspecific
taxa in the Tortula subulata complex (Pottiaceae, Bryophyta)
and their taxonomic disposition. Taxon. 56:949–952.
Cano MJ. 2008. Taxonomic revision of Hennediella Paris
(Pottiaceae, Bryophyta). Bryophytorum Bibliotheca. 64:1–
142.
Caparrós R, Lara F, Draper I, Mazimpaka V, Garilleti R. 2016.
Integrative taxonomy sheds light on an old problem: the
Ulota crispa complex (Orthotrichaceae, Musci). Botanical
Journal of the Linnean Society. 180:427–451.
Casares-Gil A. 1919. Flora Ibérica Briófitas, Hepáticas. Madrid:
Museo Nacional de Ciencias Naturales.
Cezón K, Muñoz J, Hedenäs L, Huttunen S. 2010.
Rhynchostegium confusum, a new species from the Iberian
Peninsula and its relation to R. confertum based on morphological and molecular data. Journal of Bryology. 32:1–8.
Chavoutier L, Hugonnot V. 2013. Mousses, hépatiques &
anthocérotes du départment de la Savoie (France).
Sevrier: FMBDS.
Cole TCH, Hilger HH, Goffinet B. 2019. Bryophyte phylogeny
poster (BPP). PeerJPreprints 7: e27571v1.
Cooper ED, Söderström L, Hagborg A, von Konrat MJ. 2013.
Notes on Early Land Plants Today. 38. New combinations
and synonyms in Lepidoziaceae (Marchantiophyta).
Phytotaxa 97:52–62.
Corley MFV, Crundwell AC, Düll R, Hill MO, Smith AJE. 1981.
Mosses of Europe and the Azores; an annotated list of
species, with synonyms from the recent literature.
Journal of Bryology. 11:609–689.
Cox CJ, Goffinet B, Wickett NJ, Boles SB, Shaw AJ. 2010. Moss
diversity: a molecular phylogenetic analysis of genera.
Phytotaxa. 9:175–195.
Crandall-Stotler B, Stotler RE. 2007. On the identity of Moerckia
hibernica (Hook.) Gottsche (Moerckiaceae fam. nov,
Marchantiophyta). Nova Hedwigia, Beihefte. 131:41–59.
Cros RM, Sáez L, Roselló JA. 2005. Plagiochasma appendiculatum
Lehm. & Lindenb. (Marchantiales, Aytoniaceae), a species new
to the European bryophyte flora. Journal of Bryology. 27:3–6.
Crosby MR, Buck WR. 2011. Nogopterium, a new name for the
genus Pterogonium (Musci, Leucodontaceae). Novon.
21:424–425.
Crozals A. 1903. Riccia subbifurca Warn. in litt. Revue
Bryologique. 30:62–64.
Czernyadjeva IV, Kuznetsova OI, Ignatov MS. 2017. On Pohlia
saprophila (Mielichhoferiaceae, Bryophyta). Arctoa. 26:181–
186.
Dalton NJ, Kungu EM, Long DG. 2012. The misapplication of
Hedwigia integrifolia P.Beauv. and identity of
Gymnostomum imberbe Sm. (Hedwigiaceae, Bryopsida).
Journal of Bryology. 34:59–61.
Damayanti L, Muñoz J, Wicke S, Symmank L, Shaw B, Frahm
J-P, Quandt D. 2012. Common but new: Bartramia rosamrosiae, a ‘new’ widespread species of apple mosses
(Bartramiales, Bryophytina) from the Mediterranean and
western North America. Phytotaxa. 73:37–59.
Damsholt K. 1994. On the identity of Jungermannia groenlandica Nees. Journal of the Hattori Botanical Laboratory.
75:173–178.
Damsholt K. 2002. Illustrated flora of Nordic liverworts and
hornworts. Lund: Nordic Bryological Society.
Damsholt K. 2013. The liverworts of Greenland. Lund: Nordic
Bryological Society.
Damsholt K. 2017. The complex liverwort flora of the Faeroe
Isles. Lindbergia. 40:14–38.
De Luna E. 2016. Typification, taxonomy and worldwide distribution of Braunia secunda (Hook.) Bruch, Schimp.
(Hedwigiaceae). Journal of Bryology. 38:286–294.
De Roo RT, Hedderson TA, Söderström L. 2007. Molecular
insights into the phylogeny of the leafy liverwort family
Lophoziaceae Cavers. Taxon. 56:301–314.
De Sloover JL 1959. Considérations sur la valeur spécifique de
Moerckia flotoviana (Nees) Schiffn., Dilaenacée nouvelle
pour la flore Belge. Bulletin du Jardin Botanique de l’État,
Bruxelles. 29:157–181.
Delgadillo-Moya C. 2015. Grimmia (Grimmiaceae, Bryophyta)
in the Neotropics. México City: Universidad Nacional
Autónoma de México.
Dias E, Mendes C, Shaw J. 2009. Sphagnum recurvum P.Beauv.
on Terceira, Azores, new to Macaronesia-Europe. Journal of
Bryology. 31:199–201.
Dirkse GM, Bouman AC, Losada-Lima A. 1993. Bryophytes of
the Canary Islands, an annotated checklist. Cryptogamie,
Bryologie, Lichénologie. 14:1–47.
Dirkse GM, Brugués M. 2010. Entosthodon kroonkurk
(Bryophyta: Funariaceae), a new species from the
Iberian Peninsula and Macaronesia. Journal of Bryology.
32:133–139.
Dirkse GM, Losada-Lima A, Stech M. 2016. Riccia boumanii
Dirkse, Losada & M.Stech sp. nov. (Ricciaceae,
Marchantiophyta) in the Canary Islands, the first species
of Riccia subgenus Riccia section Pilifer Volk outside
South Africa. Journal of Bryology. 38:94–102.
Dixon HN. 1912. Eucladium verbanum Nicholson and Dixon,
sp. nov. Revue Bryologique. 39:89–92.
Draper I, González-Mancebo JM, Werner O, Patiño J, Ros
RM. 2011. Phylogeographic relationships between the
mosses Exsertotheca intermedia from Macaronesian
islands and Neckera baetica from southern glacial
refugia of the Iberian Peninsula. Annales Botanici
Fennici 48:133–141.
Draper I, Hedenäs L. 2008. Sciuro-hypnum tromsoeense
(Kaurin, Arnell) Draper & Hedenäs, a distinct species
from the European mountains. Journal of Bryology.
30:271–278.
Draper I, Hedenäs L. 2009. Sciuro-hypnum dovrense (Limpr.)
Draper & Hedenäs comb. nov., a distinct Eurasian alpine
species. Cryptogamie, Bryologie. 30:289–299.
Draper I, Hedenäs L, Stech M, Patiño J, Werner O, GonzálezMancebo JM, Sim-Sim M, Lopes T, Ros RM. 2015. How
many species of Isothecium (Lembophyllaceae,
Bryophyta) are there in Macaronesia? A survey using integrative taxonomy. Botanical Journal of the Linnean Society.
177:418–438.
JOURNAL OF BRYOLOGY
Eckel P. 1998. A re-evaluation of Tortella (Musci, Pottiaceae) in
conterminous USA and Canada, with a treatment of the
European species Tortella nitida. Bulletin of the Buffalo
Society of Natural Sciences. 36:117–191.
Eckel P. 2010. Tortella bambergeri in North America and an
evaluation of its taxonomy. Bulletin of the Buffalo Society
of Natural Sciences. 39:1–10.
Eckel P. 2012. The genus Elodium (Bryophyta, Helodontiaceae):
a history and new combinations. Phytoneuron. 70:1–7.
Eckstein J, Garilleti R, Lara F. 2018. Lewinskya transcaucasica
(Orthotrichaceae, Bryopsida) sp. nov. A contribution to
the bryophyte flora of Georgia. Journal of Bryology.
40:31–38.
Eggers J. 1982. Artenliste der Moose Makaronesiens.
Cryptogamie, Bryologie, Lichénologie. 3:283–335.
Ellis LT, Afonina OM, Andriamiarisoa RL, Asthana G, Bharti R,
Aymerich P, Bambe B, Boiko M, Brugués M, Ruiz E, et al.
2018. New national and regional records, 56. Journal of
Bryology. 40:271–296.
Ellis LT, Akhoondi Darzikolaei S, Shirzadian S, Bakalin VA,
Bednarek-Ochyra H, Ochyra R, Claro D, Dulin MV, Eckel
PM, Erzberger P, et al. 2011. New national and regional
bryophyte records, 29. Journal of Bryology. 33:316–323.
Ellis LT, Aleffi M, Alegro A, Segota V, Asthana AK, Gupta R,
Singh VJ, Bakalin VA, Bednarek-Ochyra H, CykowskaMarzencka B, et al. 2016. New national and regional bryophyte records, 48. Journal of Bryology. 38:235–259.
Ellis LT, Amélio LA, Peralta DF, Bačkor M, Baisheva EZ,
Bednarek-Ochyra H, Burghardt M, Czernyadjeva IV, Kholod
SS, Potemkin AD, et al. 2019. New national and regional
bryophyte records, 59. Journal of Bryology. 41:177–194.
Ellis LT, Price MJ. 2015. Review of the type specimens of
species described by J. Hedwig in Phascum Hedw.
(Pottiaceae). Journal of Bryology. 37:23–41.
Engel JJ, Smith Merrill GL. 2004. Austral hepaticae. 35. A taxonomic
and
phylogenetic
study
of
Telaranea
(Lepidoziaceae), with a monograph of the genus in temperate Australasia and commentary on extra-Australasian
taxa. Fieldiana: Botany (n.ser.). 44:1–265.
Enroth J, Olsson S, Huttunen S, Buchbender V, Tangney R,
Stech M, Hedenäs L, Quandt D. 2019. Orthostichellaceae
fam. nov. and other novelties in pleurocarpous mosses
revealed by phylogenetic analyses. The Bryologist.
122:219–245.
Fedosov VE. 2012. Encalypta Sect. Rhabdotheca in Russia.
Arctoa. 21:101–112.
Fedosov VE, Doroshina GYa. 2018. Orthotrichales. In: Ignatov
MS, editor. Moss flora of Russia. Vol. 4. Bartramiales–
Aulacomniales. Moscow: KMK Scientific Press Ltd. (Arctoa.
27, supplement 1); p. 67–195.
Fedosov VE, Doroshina GYa, Tubanova DY, Afonina OM,
Ignatova EA. 2017. Four Orthotrichaceae species new for
Russian flora. Arctoa. 26:154–165.
Fedosov VE, Fedorova AV, Fedosov AE, Ignatov MS. 2016.
Phylogenetic inference and peristome evolution in haplolepidous mosses, focusing on Pseudoditrichaceae and
Ditrichaceae s.l. Botanical Journal of the Linnean Society.
181:139–155.
Fedosov VE, Fedorova AV, Ignatova EA. 2017a. On the taxonomic position of the genera Brachydontium Fürnr. and
Campylostelium Bruch & Schimp. (Bryophyta, Grimmiales).
Journal of Bryology. 39:161–170.
Fedosov VE, Fedorova AV, Ignatova EA. 2017b. On the two
poorly known Orthotrichum species from North Asia.
Arctoa. 26:144-153.
Fedosov VE, Fedorova AV, Ignatova EA, Ignatov MS. 2017. A
revision of the genus Seligeria (Seligeriaceae, Bryophyta)
73
in Russia inferred from molecular data. Phytotaxa.
323:27–50.
Fedosov VE, Fedorova AV, Troitsky AV, Bobrova VK, Ignatov
MS. 2016. On the systematic position of Hymenoloma
(Bryophyta). Arctoa. 25:119–130.
Fedosov VE, Ignatova EA. 2010. Orthotrichum dagestanicum
sp. nov. (Orthotrichaceae, Musci) – a new species from
Dagestan (Eastern Caucasus). Arctoa. 19:69–74.
Fedosov VE, Ignatova EA, Ivanov OV, Doroshina DY. 2010. On
the genus Entosthodon (Funariaceae, Musci) in the
Caucasus. Arctoa. 19:75–86.
Feldberg K, Groth H, Wilson R, Schäfer-Verwimp A, Heinrichs J.
2004. Cryptic speciation in Herbertus (Herbertaceae,
Jungermanniopsida): range and morphology of Herbertus
sendtneri inferred from nr ITS sequences. Plant
Systematics and Evolution. 249:247–261.
Flatberg KI. 2008. Sphagnum venustum (Bryophyta), a noticeable new species in sect. Acutifolia from Labrador, Canada.
Lindbergia. 33:2–12.
Frahm J-P. 2005. New or interesting records of bryophytes
from the Azores. Tropical Bryology. 26:45–48.
Frahm J-P. 2006. Frullania tamarisci var. azorica (Jubulaceae,
Marchantiopsida), a new taxon from the Azores. Tropical
Bryology. 27:101–105.
Frahm J-P. 2013. Are there endemic bryophyte species in the
Alps? The range of Brotherella lorentziana (Lor.) Loeske.
Archive for Bryology. 183:1–7.
Frahm J-P, Klöcker T, Schmidt R, Schröter C. 2000. Revision der
Gattung Amphidium (Musci, Dicranaceae). Tropical
Bryology. 18:173–184.
Frahm J-P, Sabovljević M. 2006. Preliminary results of the
taxonomic value of Tortula densa (Velen.) J.-P. Frahm
inferred from the Internal Transcribed Spacer (ITS) of the
nrDNA. Cryptogamie, Bryologie. 27:405–412.
Frahm J-P, Walsemann E. 1973. Nachträge zur Moosflora von
Schleswig-Holstein. Mitteilungen Arbeitsgemeinschaft
Geobotanik in Schleswig-Holstein und Hamburg. 23:1–203.
Frey W, Stech M. 2009. Marchantiophyta, Bryophyta,
Anthocerotophyta. In: Frey W, editor. Syllabus of plant
families. A. Engler’s Syllabus der Pflanzenfamilien, 13th edn,
Part 3. Bryophytes and seedless vascular plants. Stuttgart:
Gebr. Borntraeger Verlagsbuchhandlung; p. 13–263.
Furness SB, Gilbert OL 1980. The status of Thamnobryum
angustifolium (Holt) Crundw.) Journal of Bryology.
11:139–144.
Gallego MT, Cano MJ, Ros RM, Guerra J. 2002. An overview of
Sytrichia ruralis complex (Pottiaceae: Musci) in the
Mediterranean region and neighbouring areas. Botanical
Journal of the Linnean Society. 138:209–224.
Gallego MT, Hugonnot V, Cano MJ. 2018. Taxonomic resurrection of an awnless variety of Syntrichia ruralis and comparison with other European muticous taxa in this genus.
Journal of Bryology. 40:244–250.
Garcia C, Lara F, Sérgio C, Sim-Sim M, Garilleti R, Mazimpaka V.
2006. Zygodon catarinoi (Orthotrichaceae, Bryopsida), a
new epiphytic species from the Western Mediterranean
Basin. Nova Hedwigia. 82:247–256.
Gardiner A, Ignatov MS, Huttunen S, Troitsky A. 2005. On resurrection of the families Pseudoleskeaceae Schimp. and
Pylaisiaceae Schimp. (Musci, Hypnales). Taxon 54:651–663.
Geissler P. 1985. Notulae Bryofloristicae Helveticae II.
Candollea. 40:193–200.
Goffinet B, Buck WR. 2011. Physcomitridium readeri is the
correct name for Ephemerella readeri. The Bryologist.
114:545–546.
Goffinet B, Buck WR, Shaw AJ. 2009. Morphology, anatomy
and classification of the Bryophyta. In: Goffinet B, Shaw
74
N. G. HODGETTS ET AL.
AJ, editors. Bryophyte Biology, 2nd edition. Cambridge:
Cambridge University Press; p. 55–138.
Goffinet B, Budke JM, Newman LC. 2011. Micromitriaceae: a new
family of highly reduced mosses. Taxon. 60:1245–1254.
Goffinet B, Shaw AJ, Cox CJ, Wickett NJ, Boles S. 2004.
Phylogenetic inferences in the Orthotrichoideae
(Orthotrichaceae: Bryophyta) based on variation in four
loci from all genomes. Monographs in Systematic Botany
from the Missouri Botanical Garden. 98:270–289.
González-Mancebo JM, Romaguera F, Ros RM, Patiño J, Werner
O. 2008. Bryophyte flora of the Canary Islands: an updated
compilation of the species list with an analysis of distribution patterns in the context of the Macaronesian
Region. Cryptogamie, Bryologie. 29:315–357.
Gottsche CM, Lindenberg JBW, Nees CG. 1844. Synopsis hepaticarum, fasc. 1. Hamburg: Meissner; p. 1–144.
Gottsche CM, Lindenberg JBW, Nees CG. 1846. Synopsis hepaticarum, fasc. 4. Hamburg: Meissner; p. 465–624.
Grims F. 1999. Die Laubmoose Österreichs. Catalogus Florae
Austriae, II. Teil, Bryophyten (Moose), Heft 1, Musci (Laubmoose).
Wien: Österreichische Akademie der Wissenschaften.
Grolle R. 1963. Über Kurzia v. Martens. Revue Bryologique &
Lichénologique. 32:166–180.
Grolle R. 1964. Miscellanea hepaticologica 11–20. Journal of
Japanese Botany. 39:173–178.
Grolle R. 1967. Miscellanea hepaticologica 71–80. Transactions
of the British Bryological Society. 5:271–282.
Grolle R. 1973. Miscellanea hepaticologica 131–140. Herzogia.
3:75–82.
Grolle R. 1975. Miscellanea hepaticologica 151–160.
Lindbergia. 3:47–56.
Grolle R, Long DG. 2000. An annotated check-list of the
Hepaticae and Anthocerotae of Europe and Macaronesia.
Journal of Bryology. 22:103–140.
Grolle R, Persson H. 1966. Die Gattung Tylimanthus auf den
Atlantischen Inseln. Svensk Botanisk Tidskrift. 60:164–174.
Grolle R, So ML. 2003. Riccia fruticulosa O.F.Müll., 1782 and
blue Metzgeria (Marchantiophyta) in Europe. Botanical
Journal of the Linnean Society. 142:229–235
Grundmann M, Schneider H, Russell SJ, Vogel JC. 2006.
Phylogenetic relationships of the moss genus
Pleurochaete Lindb. (Bryales: Pottiaceae) based on chloroplast and nuclear genomic markers. Organisms Diversity
and Evolution. 6:33-45.
Guerra J. 1985. Orthotrichum bistratosum (Schiffn.) Guerra,
comb. & stat. nov. Anales Jardin Botánico de Madrid. 42:284.
Guerra J. 2014. Fontinalaceae. In: Guerra J, Cano MJ, Brugués M,
editors. Flora Briofítica Ibérica. Vol. V. Murcia: Universidad de
Murcia, Sociedad Española de Briología; p. 155–163.
Guerra J. 2016. Notes on the genus Eurhynchiastrum
(Bryophyta, Brachytheciaceae) in the Iberian Peninsula.
Nova Hedwigia. 102:359–365.
Guerra J. 2018. Hypnales. In: Guerra J, Cano MJ, Brugués M,
editors. Flora Briofítica Ibérica. Vol. VI. Murcia: Universidad
de Murcia, Sociedad Española de Briología; p. 365.
Guerra J, Brugués M, Cano MJ, Cros RM, editors. 2006. Flora
Briofítica Ibérica. Vol. III, Pottiaceae, Encalyptaceae,
Bartramiaceae. Murcia: Universidad de Murcia, Sociedad
Española de Briología.
Guerra J, Gallego MT, Jiménez JA, Cano MJ. 2010. Bryum Hedw.
In: Guerra, J. Brugués M, Cano MJ, Cros RM, editors. Flora
Briofítica Ibérica. Vol. IV Murcia: Universidad de Murcia &
Vitoria: Sociedad Española de Briología; p. 105–178.
Guerra J, Jiménez-Martínez, JF, Cano MJ, Alonso, M, Gallego
MT. 2019. Schistidium convergens (Grimmiaceae,
Bryophyta), a new species from southern Spain and
Morocco. Nova Hedwigia. 109:65–80.
Guerra J, Jiménez-Martínez JF, Cano MJ, Brugués M.
2008. Molecular and morphological evidence on
the status of Bryum mildeanum Jur. and records
revision in the Iberian Peninsula. Journal of Bryology.
30:297–302.
Guerra J, Jiménez-Martínez JF, Cano MJ, Jiménez-Fernández
JA. 2011. A contribution to the phylogenetic study of
Mielichhoferiaceae-Mniaceae (Bryophyta) based on molecular sequence data. Nova Hedwigia. 93:47–56.
Guerra J, Jiménez-Martínez, JF, Jiménez, JA. 2010. Neckera
baetica sp. nov. (Neckeraceae, Bryophyta) from southern
Spain, based on morphological and molecular data. Nova
Hedwigia. 91:255–263.
Guerra J, Martínez M, Jiménez JA, Cano MJ, Gallego MT. 2020.
A new species of moss emerges from molecular and morphological data: Schistidium memnonium sp. nov.
(Grimmiaceae, Bryophyta). Plant Biosystems. https://doi.
org/10.1080/11263504.2020.1762789.
Guerra J, Ríos D, Gallego MT. 2014. A revision of the genus
Rhynchostegiella (Brachytheciaceae) in the Iberian
Peninsula and Balearic Islands. Phytotaxa. 183:193–200.
Hampe E. 1872. Symbolae ad floram Brasiliae centralis cognoscendam. Musci frondosi. Vidensk. Meddel. Dansk
Naturhist. Foren. Kjøbenhavn 4:36–59
Hanssen L, Såstad SM, Flatberg KI. 2000. Population structure
and taxonomy of Sphagnum cuspidatum and S. viride. The
Bryologist. 103:93–103.
Hassel K, Kyrkjeeide MO, Yousefi N, Prestø T, Stenøien HK,
Shaw JA, Flatberg KI. 2018. Sphagnum divinum (sp. nov.)
and S. medium Limpr. and their relationship to
S. magellanicum Brid. Journal of Bryology. 40:197–222.
Hattori S. 1972. Frullania tamarisci complex and the species
concept. Journal of the Hattori Botanical Laboratory.
35:202–251.
Hedderson TA, Blockeel TL. 2006. Oncophorus dendrophilus, a
new moss species from Cyprus and Crete. Journal of
Bryology. 28:357–359.
Hedderson TA, Murray DJ, Cox CJ, Nowell TL. 2004.
Phylogenetic relationships of haplolepidous mosses
(Dicranidae) inferred from rps4 gene sequences.
Systematic Botany. 29:29–41.
Hedderson TA, Zander RH. 2008. Chenia ruigtevleia
(Pottiaceae), a new moss species from the Western Cape
Province of South Africa. The Bryologist. 111:496–500.
Hedenäs L. 1994. The Hedwigia ciliata complex in Sweden,
with notes on the occurrence of the taxa in
Fennoscandia. Journal of Bryology. 18:139–157.
Hedenäs L. 2006. Additional insights into the phylogeny of
Calliergon, Loeskypnum, Straminergon and Warnstorfia
(Bryophyta: Calliergonaceae). Journal of the Hattori
Botanical Laboratory. 100:125–134.
Hedenäs L. 2010. ITS distinguishes Palustriella falcata from
P. commutata and P. decipiens, but not from
P. pluristratosa (Bryophyta: Amblystegiceae). Nova
Hedwigia, Bieheft. 138:51–60.
Hedenäs L. 2012. Global phylogeography in Sanionia uncinata
(Amblystegiaceae: Bryophyta). Botanical Journal of the
Linnean Society. 168:19–42.
Hedenäs L. 2017a. Scandinavian Oncophorus (Bryopsida,
Oncophoraceae): species, cryptic species, and intraspecific
variation. European Journal of Taxonomy. 315:1–34.
Hedenäs L. 2017b. Brachythecium tauriscorum Molendo, a
widely distributed Arctic-alpine species. Journal of
Bryology. 39:201–203.
Hedenäs L. 2018. Oncophorus demetrii, a fifth Scandinavian
species of Oncophorus (Musci) possible to recognize by
morphology. Lindbergia. 41:1–9.
JOURNAL OF BRYOLOGY
Hedenäs L. 2020. Disentangling Scandinavian species hidden
within Meesia uliginosa Hedw. s.l. (Bryophyta, Meesiaceae).
Lindbergia. 42(1):1–14. DOI: 10.25227/linbg.01125
Hedenäs L, Désamoré A, Laenen B, Papp B, Quandt D,
González-Mancebo JM, Patiño J, Vanderpoorten A,
Stech M. 2014. Three species for the price of one
within the moss Homalothecium sericeum s.l. Taxon.
63:249–257.
Hedenäs L, Heinrichs J, Gallego MT. 2019. The Scandinavian
Syntrichia ruralis complex (Musci. Pottiaceae): a chaos
of diversification. Plant Systematics and Evolution.
DOI: 10.1007/s00606-019-01596-0
Hedenäs L, Rosborg C. 2008. Pseudocalliergon is nested within
Drepanocladus (Bryophyta: Amblystegiaceae). Lindbergia.
33:67–74.
Heinrichs J, Dong S, Schäfer-Verwimp A, Pócs T, Feldberg K,
Czumaj A, Schmidt AR, Reitner J, Renner MAM, Hentschel
J, et al. 2013. Molecular phylogeny of the leafy liverwort
Lejeunea (Porellales): evidence for a Neotropical origin,
uneven distribution of sexual systems and insufficient taxonomy. PLOS one. 8(12;E82547):1–14.
Heinrichs J, Hentschel J, Bombosch A, Fiebig A, Reise J,
Edelmann M, Kreier HP, Schäfer-Verwimp A, Caspari S,
Schmidt AR, et al. 2010. One species or at least eight?
Delimitation and distribution of Frullania tamarisci (L.)
Dumort. s. l. (Jungermanniopsida, Porellales) inferred
from nuclear and chloroplast DNA markers. Molecular
Phylogenetics and Evolution. 56:1105–1114.
Heinrichs J, Kreier HP, Feldberg K, Schmidt AR, Zhu RL, Shaw
B, Shaw AJ, Wiessman V. 2011. Formalizing morphologically cryptic biological entities: new insights from DNA taxonomy, hybridization, and biogeography in the leafy
liverwort
Porella
platyphylla
(Jungermanniopsida,
Porellales). American Journal of Botany. 98:1252–1262.
Heinrichs J, Pröschold T, Renker C, Groth H, Rycroft DS. 2002.
Plagiochila virginica A.Evans rather than P. dubia Lindenb. &
Gottsche occurs in Macaronesia; placement in sect.
Contigua Carl is supported by ITS sequences of nuclear
ribosomal DNA. Plant Systematics and Evolution.
230:221–230.
Heinrichs J, Rycroft DS, Groth H, Cole WJ. 2002. Morphological
and phytochemical studies of Plagiochila papillifolia Steph.,
a Neotropical liverwort new to Europe. Journal of Bryology.
24:119–126.
Hentschel J, von Konrat M, Söderström L, Hagborg A, Larraín J,
Sukkharak P, Uribe J, Zhang L. 2015. Notes on Early Land
Plants Today. 72. Infrageneric classification and new combinations, new names, new synonyms in Frullania.
Phytotaxa. 220:127–142.
Hernández-Maqueda R, Quandt D, Muñoz J. 2008. Testing
reticulation and adaptive convergence in the
Grimmiaceae (Bryophyta). Taxon. 57:500–510.
Hill MO, Bell N, Bruggeman-Nannenga MA, Brugués M, Cano
MJ, Enroth J, Flatberg KI, Frahm J-P, Gallego MT, Garilleti
R, et al. 2006. An annotated checklist of the mosses of
Europe and Macaronesia. Journal of Bryology. 28:198–267.
Hill MO, Blackstock TH, Long DG, Rothero GP. 2008. A checklist and census catalogue of British and Irish bryophytes.
Middlewich: British Bryological Society.
Hodgetts NG 2003. Some synonyms of Herbertus dicranus
(Taylor ex Gottsche et al.) Trevis. Journal of Bryology.
25:138–140.
Hodgetts NG. 2015. Checklist and country status of European
bryophytes – towards a new Red List for Europe. Irish
Wildlife Manuals 84. Dublin: National Parks & Wildlife
Service, Department of Arts, Heritage and the Gaeltacht,
Ireland.
75
Hodgetts NG, Blockeel TL. 1992. Thamnobryum cataractarum,
a new species from Yorkshire, with observations on
T. angustifolium and T. fernandesii. Journal of Bryology.
17:251–262.
Hodgetts N, Cálix M, Englefield E, Fettes N, García Criado M,
Patin L, Nieto A, Bergamini A, Bisang I, Baisheva E, et al.
2019. A miniature world in decline: European Red List of
mosses, liverworts and hornworts. Brussels: International
Union for the Conservation of Nature.
Hodgetts NG, Vanderpoorten A. 2018. Isothecium myosuroides
var. brachythecioides (Dixon) Braithw. reinstated as a
species, I. interludens Stirt. Journal of Bryology. 40:316–323.
Holyoak DT. 2003. A taxonomic review of some British coastal
species of the Bryum bicolor complex, with a description of
Bryum dyffrynense sp. nov. Journal of Bryology. 25:107–113.
Holyoak DT. 2009. Bryum apiculatum (Bryaceae) in East
Cornwall: a moss new to Europe. Journal of Bryology.
31:117–120.
Holyoak DT. 2010. Notes on taxonomy of some European
species of Ephemerum (Bryopsida: Pottiaceae). Journal of
Bryology. 32:122–132.
Holyoak DT, Hedenäs L. 2006. Morphological, ecological and
molecular studies of the intergrading taxa Bryum neodamense and B. pseudotriquetrum (Bryopsida: Bryaceae).
Journal of Bryology. 28:299–311.
Holyoak DT, Köckinger H. 2010. A taxonomic revision of some
European and Asian bulbiferous species of Anomobryum
(Bryophyta: Bryaceae). Journal of Bryology. 32:153–169.
Holyoak DT, Lockhart N. 2009. Australasian bryophytes
introduced to South Kerry with tree ferns. Field Bryology.
98:3–7.
Holyoak DT, Pedersen N. 2007. Conflicting molecular and
morphological evidence of evolution within the Bryaceae
(Bryopsida) and its implications for generic taxonomy.
Journal of Bryology. 29:111–124.
Hooper EJ, Duckett JG, Cuming AC, Kunin WE, Pressel S. 2010.
Ephemerella readeri Müll.Hal. (Physcomitrella readeri
(Müll.Hal.) I.G.Stone & G.A.M.Scott, Funariidae, Bryophyta):
a genus and species new to Europe. Journal of Bryology.
32:256–264.
Hugonnot V. 2010a. Mousses & hépatiques de Païolive
(Ardèche & Gard, France). Bulletin de la Société
Botanique du Centre-Ouest, nouvelle série, numéro
special. 34:1–293.
Hugonnot V. 2010b. Towards an improved understanding of
Riccia ciliata Hoffm. (Marchantiopsida: Ricciaceae). Journal
of Bryology. 32:300–303.
Hugonnot V. 2015. Riccia melitensis Mass. (Marchantiophyta:
Ricciaceae), an endemic species of the Maltese archipelago? Phytotaxa. 222:238–240.
Hugonnot V. 2018. Bryophyte flora and vegetation of serpentine sites in the French Massif Central. Plant Biosystems.
DOI: 10.1080/11263504.2018.1527789.
Hugonnot V. 2019. New records for the bryophyte flora of
Corsica. Cryptogamie, Bryologie. 40:153–158.
Hugonnot V, Hedenäs L. 2015. Arvernella microclada
Hugonnot & Hedenäs (Amblystegiaceae), a new minute
species from France, requiring a separate genus. Journal
of Bryology. 37:184–191.
Hugonnot V, Porley RD, Ignatov MS. 2018. A taxonomic conundrum resolved: the transfer of Grimmia horrida to
Coscinodon based on sporophyte discovery in France, with
support from molecular data. The Bryologist. 121:520–528.
Hugonnot V, Schill DB. 2006. Mannia californica (Gottsche ex
Underw.) L.C.Wheeler (Aytoniaceae, Marchantiales) in
Ardèche (France), new to Europe. Cryptogamie,
Bryologie. 27:181–189.
76
N. G. HODGETTS ET AL.
Hutsemékers V, Vieira CC, Ros RM, Huttunen S, Vanderpoorten
A. 2012. Morphology informed by phylogeny reveals unexpected patterns of species differentiation in the aquatic
moss
Rhynchostegium
riparioides
s.l.
Molecular
Phylogenetics and Evolution. 62:748–755.
Huttunen S, Bell N, Bobrova VK, Buchbender V, Buck WR, Cox
CJ, Goffinet B, Hedenäs L, Ho BC, Ignatov MS, et al. 2012.
Disentangling knots of rapid evolution: origin and diversification of the moss order Hypnales. Journal of Bryology.
34:187–211.
Huttunen S, Ignatov MS. 2010. Evolution and taxonomy of
aquatic species in the genus Rhynchostegium
(Brachytheciaceae, Bryophyta). Taxon. 59:791–808.
Huttunen S, Kuznetsova O, Li M, Wang YF, Ignatov MS. 2015.
East Asian origin and Northern Hemisphere diversification
in the Brachythecium novae-angliae complex (Bryophyta).
Taxon. 64:911–930.
Ignatov MS, Afonina OM, Ignatova EA. 2006. Check-list of
mosses of East Europe and North Asia. Arctoa. 15:1–130.
Ignatov MS, Afonina OM, Kuznetsova OI, Ignatova EA. 2012.
The
genus
Leptopterigynandrum
(Taxiphyllaceae,
Bryophyta) in Russia. Arctoa. 21:207–220.
Ignatov MS, Bersanova AS, Zharzinov ZS, Ignatova EA. 2005.
Leptodontium (Pottiaceae, Bryophyta), a new genus for
Caucasus. Arctoa. 14:35–38.
Ignatov MS, Fedorova AV, Fedosov VE. 2019. On the taxonomy of Anomodontaceae and Heterocladium (Bryophyta).
Arctoa. 28:75–102.
Ignatov MS, Fedosov VE, Ignatova EA, Doroshina DY, Zolotov
VI. 2010. Moss flora of Gunib area in Dagestan, the Eastern
Caucasus. Arctoa. 19:87–96.
Ignatov MS, Gardiner AA, Bobrova VK, Milyutina IA, Huttunen
S, Troitsky AV. 2007. On the relationships of mosses of the
order Hypnales, with special reference to taxa traditionally
classified in Leskeaceae. In: Newton AE, Tangney R, editors.
Pleurocarpous mosses: systematics and evolution. Boca
Raton, CRC Press; p. 177–213.
Ignatov MS, Huttunen S, Li M, Wang Y-F. 2015. How to know
Myuroclada longiramea (Brachytheciaceae, Bryophyta).
Arctoa. 24:497–502.
Ignatov MS, Ignatova EA. 2004. Moss flora of the Middle
European Russia. Vol. 2: Fontinalaceae–Amblystegiaceae.
Arctoa. 11 (Supplement 2):609–960.
Ignatov MS, Ignatova EA, Kuznetsova OI, Fedosov VE. 2019.
On moss genera Hylocomiadelphus Ochyra & Stebel and
Rhytidiadelphus (Limpr.) Warnst. Acta Musei Silesiae,
Scientiae Naturales. 68:123–134.
Ignatov MS, Ivanova EI, Ignatova EA, Krivoshapkin KK. 2001.
On the moss flora of Ust-Maya District (Republic Sakha/
Yakutia, East Siberia). Arctoa. 10:165–184.
Ignatov MS, Milyutina IA. 2007. On Sciuro-hypnum oedipodium
and S. curtum (Brachytheciaceae, Bryophyta). Arctoa.
16:47–61.
Ignatov MS, Milyutina IA. 2010. The genus Brachythecium
(Brachytheciaceae, Musci) in Russia: comments on
species and key for identification. Arctoa. 19:1–30.
Ignatov MS, Porley RD, Ochyra R, Kuznetsova OI, Ignatova EA.
2018. Coscinodon monchiquensis R.D.Porley, Ochyra &
Ignatova (Grimmiaceae), a new species from the
Algarve, southern Portugal. Journal of Bryology. 40:125–
136.
Ignatov MS, Smith Merrill GL. 1995. Bryophytes of Altai
Mountains. VI. The family Polytrichaceae (Musci). Arctoa.
5:61–97.
Ignatova EA, Blom HH, Goryunov DV, Milyutina IA. 2010. On
the genus Schistidium (Grimmiaceae, Musci) in Russia.
Arctoa. 19:195–233.
Ignatova EA, Fedorova AV, Kuznetsova OI, Ignatov MS. 2019.
Taxonomy of the Plagiothecium laetum complex
(Plagiotheciaceae, Bryophyta) in Russia. Arctoa. 28:28–45.
Ignatova EA, Ignatov MS, Milyutina IA. 2010. A revision of the
genus Lindbergia (Leskeaceae, Bryophyta) in Russia. Arctoa.
19:97–116.
Ignatova EA, Kuznetsova OI, Fedosov VE, Ignatov MS. 2016.
On the genus Hedwigia (Hedwigiaceae, Bryophyta) in
Russia. Arctoa. 25:241–277.
Ignatova EA, Kuznetsova OI, Ignatov MS, Köckinger H. 2012.
The
genera
Oxystegus
and
Pseudosymblepharis
(Pottiaceae, Bryophyta) in the Caucasus. Arctoa. 21:173–
180.
Ignatova EA, Kuznetsova OI, Köckinger H, Hastings R. 2008. A
preliminary study of Coscinodon (Grimmiaceae, Musci) in
Eurasia based on morphology and DNA sequence data.
Arctoa. 17:1–18.
Ignatova EA, Kuznetsova OI, Milyutina IA, Fedosov VE, Ignatov
MS. 2017. The genus Fabronia (Fabroniaceae, Bryophyta) in
Russia. Arctoa. 26:11–34.
Ignatova EA, Tubanova DY, Tumurova OD, Goryunov DV,
Kuznetsova OI. 2015. When the plant size matters: a new
semi-cryptic species of Dicranum from Russia. Arctoa.
24:471–488.
Inoue H. 1969. Contributions to the knowledge of
Plagiochilaceae of southern Asia. XII. Typification of
Plagiochila javanica (Swartz) Dum. Journal of the Hattori
Botanical Laboratory. 32:50–56.
Inoue Y, Tsubota H. 2014. On the systematic position of the
genus Timmiella (Dicranidae, Bryopsida) and its allied
genera, with the description of a new family
Timmiellaceae. Phytotaxa. 181:151–162.
Inoue Y, Tsubota H. 2016. Systematics of the family Pottiaceae
(Bryophyta) with special reference to the familial and subfamilial circumscriptions. Hikobia. 17:117–129.
Ivanova EI, Bell NE, Kuznetsova OI. 2015. The genus
Polytrichum Sect. Aporotheca (Polytrichaceae) in Russia.
Arctoa. 24:67–78.
Ivanova EI, Bell NE, Kuznetsova OI, Lee H, Park M, Ignatov M.
2014. The genus Polytrichastrum (Polytrichaceae) in Russia.
Arctoa. 23:164–184.
Iwatsuki Z. 1959. Bryological miscellanies, IX. A note on
Japanese Orthotrichum. Journal of the Hattori Botanical
Laboratory. 21:240.
Jiménez JA. 2006. Taxonomic revision of the genus
Didymodon Hedw. (Pottiaceae, Bryophyta) in Europe,
North Africa and Southeast and Central Asia. Journal of
the Hattori Botanical Laboratory. 100:211–292.
Jovet-Ast S. 1986. Les Riccia de la région Méditerranéenne.
Cryptogamie, Bryologie Lichénologie. 7:287–431.
Jovet-Ast S. 2000. Documents pour la connaissance des
Riccia Australiens (Hépatiques, Marchantiales) – nouvelles récoltes. Taxons nouveaux. Commentaires morphologiques & écologiques. Cryptogamie, Bryologie.
21:289–343.
Juslén A. 2006. Revision of Asian Herbertus (Herbertaceae,
Marchantiophyta). Annales Botanici Fennici. 43:409–
436.
Kanda H. 1976 (1977). A revision of the family
Amblystegiaceae of Japan II. Journal of Science of the
Hiroshima University, Series B, Division 2, Botany. 16:47–
119.
Kiebacher T, Lüth M. 2017. Orthotrichum dentatum
T.Kiebacher, Lüth sp. nov. (Orthotrichaceae). Journal of
Bryology. 39:46–56.
Kindberg NC. 1893. Notes on Canadian bryology. Ottowa
Naturalist. 7:17–23.
JOURNAL OF BRYOLOGY
Kindberg NC. 1897. European and N. American Bryineae
(mosses). Linköping: Linköpings Lithografiska Aktiebolag.
Köckinger, H. 2016. Rediscovery and redescription of the
enigmatic Radula visianica (Porellales, Marchantiophyta).
Herzogia. 29:625–634.
Köckinger H. 2017. Die Horn- und Lebermoose Österreichs
(Anthocerotophyta und Marchantiophyta). Catalogus
Florae Austriae. II(2):1–382.
Köckinger H, Bednarek-Ochyra H, Ochyra R. 2007.
Bucklandiella nivalis (Grimmiaceae), a new moss from the
Alps of Austria. The Bryologist. 110:92–99.
Köckinger H, Hedenäs L. 2017. A farewell to Tortella bambergeri (Pottiaceae) as understood over the last decades.
Journal of Bryology. 39:213–225.
Köckinger H, Holyoak DT, Suanjak M. 2013. Bryum austriacum
Köckinger, Holyoak & Suanjak, a new bulbiferous species
from the Alps (Bryaceae, Bryopsida). Journal of Bryology.
35:57–61.
Köckinger H, Kučera J. 2007. Barbula amplexifolia (Mitt.)
A.Jaeger in Europe. Journal of Bryology. 29:33–40.
Köckinger H, Kučera J. 2011. Hymenostylium xerophilum sp.
nov., and H. gracillimum, comb. nov., two neglected
European mosses and their molecular affinities. Journal
of Bryology. 33:195–209.
Köckinger H, Kučera J. 2016. Brachythecium funkii Schimp. and
B. japygum (Głow.) Köckinger & Jan Kučera comb. nov., two
alpine species hitherto included in B. cirrosum (Schwägr.)
Schimp. Journal of Bryology. 38:267–285.
Köckinger H, Kučera J, Hofmann H, Müller N, Amann G. 2012.
Barbula consanguinea discovered in Switzerland and
Austria, with a revision of former European records of
B. indica. Herzogia. 25:61–70.
Köckinger H, Lüth M, Werner O, Ros RM. 2018. Tortella mediterranea (Pottiaceae), a new species from southern
Europe, its molecular affinities, and taxonomic notes on
T. nitida. The Bryologist. 121:560–570.
Köckinger H, Suanjak M, Schriebl A, Schröck C. 2008. Die
Moose Kärntens. Klagenfurt: Natur Kärnten, Sonderreihe
des Naturwissenschaftlichen Vereins für Kärnten.
Köckinger H, van Melick H. 2007. Didymodon maschalogenus
(Pottiaceae), a novelty in the European moss flora, reported
from the Austrian Alps and southern Norway. Lindbergia.
32:62–68.
Köckinger H, Werner O, Ros RM. 2010. A new taxonomic
approach to the genus Oxystegus (Pottiaceae, Bryophyta)
in Europe based on molecular data. Nova Hedwigia,
Beiheft. 138:31–49.
Konstantinova NA. 2000a. [Distribution patterns of the north
holarctic hepatics]. Arctoa. 9:29–94.
Konstantinova NA. 2000b. Redkie pechenochniki (Hepaticae)
Murmansko oblasti i podhody k ih ohrane [Rare liverworts
(Hepaticae) of Murmansk province and some approaches
to their protection]. Botanicheskii zhurnal. Moscow,
Leningrad. 85(10):122–135. Russian.
Konstantinova NA, Akatova TV, Savchenko AN. 2009. Hepatics
of Caucasian State Nature Reserve (Western Caucasus,
Russia). Arctoa 18:121–134.
Konstantinova NA, Bakalin VA, Andreeva EN, Bezgodov AG,
Borovichev EA, Dulin MA, Mamontov YS. 2009. Checklist
of liverworts (Marchantiophyta) of Russia. Arctoa. 18:1–
64.
Konstantinova NA, Doroshina GYa. 2011. New liverwort
records from Republic of Kalmykia. 1. Arctoa. 20:256.
Konstantinova NA, Lavrinenko OV. 2002. [On the flora of
hepaticae of the Nenets autonomous area (Archangelsk
region, north-eastern European Russia)]. Botanicheskii
zhurnal. Moscow, Leningrad 87(9):43–50. Russian.
77
Konstantinova NA, Mamontov YS, Vilnet AA. 2020. Frullania
stylifera (R.M.Schust.) R.M.Schust. (Hepaticae), new to
Eurasia. Journal of Bryology. 42: https://doi.org/10.1080/
03736687.2020.1722511
Konstantinova NA, Potemkin AD. 1994. Studies on Scapania
sphaerifera (Hepaticae). Annales Botanici Fennici. 31:121–126.
Konstantinova NA, Potemkin AD. 1996. Liverworts of the
Russian arctic: an annotated check-list and bibliography.
Arctoa. 6:125–150.
Konstantinova NA, Potemkin AD, Schljakov RN. 1992.
Checklist of the Hepaticae and Anthocerotae of the
former USSR. Arctoa. 1:87–127.
Konstantinova NA, Savchenko AN. 2013. Russian Caucasus
distribution of fourteen hepatics rare and threatened in
Europe. Polish Botanical Journal. 58:573–584.
Konstantinova NA, Savchenko AN. 2018. Contribution to the
bryophyte flora of Svalbard: hepatics of Barents Island.
Folia Cryptogamica Estonica 55:105–115.
Konstantinova NA, Vilnet AA. 2011. Jubula hutchinsiae subsp.
caucasica subsp. nov. (Jubulaceae, Marchantiophyta) – a
new taxon from the western Caucasus. Arctoa. 20:227–238.
Konstantinova NA, Vilnet AA. 2016. A new species of the
genus Jungermannia (Jungermanniales, Marchantiophyta)
from the Caucasus with notes on taxa delimitation and taxonomy of Jungermannia s. str. Phytotaxa. 255:227–239.
Koponen T. 2014. Bryophyte flora of Hunan Province, China.
18. Mniaceae subfam. Mnioideae (Musci). Acta
Bryolichenologica Asiatica. 5:39–72.
Koponen T. 2017. Lähdelehväsammal (Rhizomnium dentatum), jolla oli hampainen lehtilaita, ja sen mahdollinen
jälkeläinen, R. punctatum var. hermanperssonii, var. nov.
Bryobrotherella. 20:62–66.
Koponen T, Ignatova EA. 2018. Mniaceae. In: Ignatov MS,
editor. Moss Flora of Russia. Vol. 4. BartramialesAulacomniales. Moscow: KMK Scientific Press Ltd. (Arctoa.
27, supplement 1); p. 439–517.
Koponen T, Isoviita P. 2010. Philonotis capillaris Lindb. and
P. arnellii Husn.; one moss, two names. Cryptogamie,
Bryologie. 31:75–94.
Koponen T, Sun Y. 2017. Preliminary study on phylogenetic
position and delimitation of the ciliate arthrodontous
genera of the moss family Mniaceae. Journal of Bryology.
39:23–38.
Košnar J, Kolář F. 2009. A taxonomic study of selected
European taxa of the Tortula muralis (Pottiaceae, Musci)
complex: variation in morphology and ploidy level.
Preslia. 81:399–421.
Kramer W. 1980. Tortula Hedw. sect. Rurales De Not.
(Pottiaceae, Musci) in der östlichen Holarktis.
Bryophytorum Bibliotheca. 21:1–165.
Kučera J, Blockeel TL, Erzberger P, Papp B, Soldán Z, Vellak K,
Werner O, Ros RM. 2018. The Didymodon tophaceus
complex (Pottiaceae, Bryophyta) revisited: new data
support the subspecific rank of currently recognized
species. Cryptogamie, Bryologie. 39:241–257.
Kučera J, Košnar J, Werner O. 2013. Partial generic revision of
Barbula (Musci: Pottiaceae): re-establishment of
Hydrogonium and Streblotrichum, and the new genus
Gymnobarbula. Taxon. 62:21–39.
Kučera J, Kuznetsova OI, Manukjanová A, Ignatov MS. 2019. A
phylogenetic revision of the genus Hypnum. Towards completion. Taxon. 68:1-33.
Kučera J, Váňa J, Hradílek Z. 2012. Bryophyte flora of the
Czech Republic: updated checklist and Red List and a
brief analysis. Preslia. 84:813–850.
Kürschner H, Erdağ A. 2005. Bryophytes of Turkey: an
annotated reference list of the species with synonyms
78
N. G. HODGETTS ET AL.
from the recent literature and an annotated list of
Turkish bryological literature. Turkish Journal of Botany.
29:95–154.
Kyrkjeeide MO, Hassel, K, Stenøien HK, Prestø T, Boström E,
Shaw AJ, Flatberg KI. 2015. The dark morph of Sphagnum
fuscum (Schimp.) H.Klinggr. in Europe is conspecific with
the North American S. beothuk. Journal of Bryology.
37:251–266.
Laine J, Flatberg KI, Harju P, Timonen T, Minkkinen K, Laine A,
Tuittila ES, Vasander H. 2018. Sphagnum mosses – the stars
of European mires. Helsinki: University of Helsinki.
Lamarck, JB. 1815. Flore Française, ou, descriptions succinctes
de toutes les plantes qui croissent naturellement en
France. Paris: Desray.
Lang AS, Bocksberger G, Stech M. 2015. Phylogeny and
species delimitations in European Dicranum (Dicranaceae,
Bryophyta) inferred from nuclear and plastid DNA.
Molecular Phylogenetics and Evolution. 92:217–225.
Lara F, Garilleti R. 2014. Orthotrichum Hedw. In: Guerra J, Cano
MJ, Brugués M, editors. Flora Briofítica Ibérica, Vol V.
Murcia: Universidad de Murcia, Sociedad Española de
Briología; p. 50–135.
Lara F, Garilleti R, Draper I, Mazimpaka V. 2018. Lewinskya
lamyana sp. nov. (Orthotrichaceae, Bryopsida), a distinct
moss from an exceptional habitat in the southern Iberian
Peninsula. Cryptogamie, Bryologie. 39:259–270.
Lara F, Garilleti R, Goffinet B, Draper I, Medina R, Vigalondo B,
Mazimpaka V. 2016. Lewinskya, a new genus to accommodate the phaneroporous and monoicous taxa of
Orthotrichum (Bryophyta, Orthotrichaceae). Cryptogamie,
Bryologie. 37:361–382.
Lara F, Mazimpaka V, Estébanez B, Garilleti R. 2009.
Orthotrichum consobrinum Cardot in western Europe and
south-western Asia. Journal of Bryology. 31:80–85.
Larraín J, Quandt D, Stech M, Muñoz J. 2013. Lumping or splitting? The case of Racomitrium (Bryophytina: Grimmiaceae).
Taxon. 62:1117–1132.
Ligrone R, Duckett JG. 2005. Cyathodium Kunze
(Cyathodiaceae, Marchantiophyta), genere e famiglia
nuovi per l’Europa, nella Valle delle Ferriere (Amalfi,
Salerno). Informatore Botanico Italiano. 37:84–85.
Limpricht KG. 1895. 1. Die Laubmoose Deutschlands,
Oesterreichs und der Schweiz. II. Abteilung: Bryinae. In:
Rabenhorst L. Kryptogamen-Flora von Deutschland,
Oesterreich und der Schweiz. 4. Leipzig: Eduard Kummer.
Limpricht G. 1897. Drei neuen Laubmoose. Jahresbericht der
Schlesischen Gesellschaft für Vaterländische Cultur. 74
(2b):1–5.
Lindberg SO. 1877. Monographia Metzgeriae. Acta Societatis
pro Fauna & Flora Fennica. 1:1–49.
Loeske L. 1929. Die Laubmoos Europas II Funariaceae.
Repertorium Speciarum Novarum Regni Vegetabilis,
Sonderbeiheft B. 3(2):1–71.
Long DG 2006. Revision of the genus Asterella P. Beauv. in
Eurasia. Bryophytorum Bibliotheca. 63:1–299.
Long DG, Paton JA, Squirrell J, Woodhead M, Hollingsworth
PM. 2006. Morphological, ecological and genetic evidence
for distinguishing Anastrophyllum joergensenii Schiffn. and
A. alpinum Steph. (Jungermanniopsida: Lophoziaceae).
Journal of Bryology. 28:108–117.
Lönnell N, Hassel K. 2018. New combinations within the
genus Sphagnum (Sphagnaceae, Bryophyta). Phytotaxa.
369:57–58.
Maier E. 2002. Grimmia dissimulata E.Maier sp. nova, and
the taxonomic position of Grimmia trichophylla var. meridionalis Müll.Hal. (Musci, Grimmiaceae). Candollea.
56:281–300.
Maier E. 2010. The genus Grimmia Hedw. (Grimmiaceae,
Bryophyta) — a morphological anatomical study.
Boissiera. 63:5–377.
Majestyk P. 2011. A taxonomic treatment of Daltonia (Musci:
Daltoniaceae) in the Americas. Journal of the Botanical
Research Institute of Texas. 5:553–575.
Mamontov YS, Konstantinova NA, Vilnet AA. 2018. One more
species in the genus Jungermannia (Marchantiophyta:
Jungermanniaceae). Bryophyte Diversity and Evolution.
40:79–90.
Mamontov YS, Potemkin AD, Tubanova DY, Sofronova EV.
2018. [Liverworts of the Dzherginsky Reserve (Republic
of Buryatia)]. Novosti sistematiki nizshikh rastenii.
52:483–504.
Manukjanová A, Kouteckí P, Štechová T, Kučera J. 2019.
Insights into the distribution patterns, habitat and morphologic differentiation of cryptic species of the moss
Hamatocaulis vernicosus in the Czech Republic. Herzogia.
32:183–199.
Marka J, Blockeel TL, Long DG, Papp B. 2018. Bryophytes new
to Albania from the British Bryological Society field
meeting in 2014. Journal of Bryology. 40:163–172.
Mastracci M. 2003. Thamnobryum neckeroides (Bryopsida:
Neckeraceae): lectotypification, synonymies, diagnostic
characters, habitat and distribution. Journal of Bryology.
25:115–120.
Masuzaki H, Shimamura M, Furuki T, Tsubota H, Yamaguchi T,
Majid HMA, Deguchi H. 2010. Systematic position of the
enigmatic
liverwort
Mizutania
(Mizutaniaceae,
Marchantiophyta) inferred from molecular phylogenetic
analyses. Taxon. 59:448–458.
Mazimpaka V, Lara F. 2014. Codonoblepharon Schwägr. In:
Guerra J, Cano MJ, Brugués M, editors. Flora Briofítica
Ibérica, Vol. V. Murcia: Universidad de Murcia, Sociedad
Española de Briología; p. 27–30.
Medina R, Johnson MG, Liu Y, Wickett NJ, Shaw AJ, Goffinet B.
2019. Phylogenetic delineation of Physcomitrium
(Bryophyta: Funariaceae) based on targeted sequencing
of nuclear exons and their flanking regions rejects the
retention of Physcomitrella, Physcomitridium and
Aphanorrhegma. Journal of Systematics and Evolution.
57:404–417.
Medina R, Lara F, Goffinet B, Garilleti R, Mazimpaka V. 2012.
Integrative taxonomy successfully resolves the pseudocryptic complex of the disjunct epiphytic moss
Orthotrichum consimile s.l. (Orthotrichaceae). Taxon.
61:1180–1198.
Medina R, Lara F, Goffinet B, Garilleti R, Mazimpaka V. 2013.
Unnoticed diversity within the disjunct moss
Orthotrichum tenellum s.l. validated by morphological and
molecular approaches. Taxon. 62:1133–1152.
Meinunger L, Schröder W. 2007. Verbreitungsatlas der Moose
Deutschlands. Regensburg: O. Dürhammer für die
Regensburgische Botanische Gesellschaft.
Mitten W 1859. Musci Indiae Orientalis, an enumeration of the
mosses of the East Indies. Journal of the Proceedings of the
Linnean Society, Botany. 1:1–96.
Mitten W. 1864. Contributions to the Cryptogamic Flora of the
Atlantic Islands. Journal of the Linnean Society. Botany.
8:1–10.
Müller C. 1849. Synopsis Muscorum Frondosorum omnium
hucusque Cognitorum. Vol. 1. Berolini [Berlin]: A. Foerstner.
Müller C. 1858. Synopsis Muscorum Frondosorum omnium
hucusque Cognitorum. Berlin: A. Foerstner.
Müller F. 2007. Zum Vorkommen von Clasmatodon parvulus
(Fabroniaceae, Bryopsida) in Deutschland. Herzogia.
20:293–298.
JOURNAL OF BRYOLOGY
Müller F. 2014. Bartramia aprica - the correct name for the
Mediterranean and western North American species historically recognized as “Bartramia stricta”. Herzogia.
27:211–214.
Müller K. 1953. Hepatikologische Notizen. Revue Bryologique
& Lichénologique. 22:131–140.
Müller K. 1954. Die Lebermoose Europas (Dr. L. Rabenhorst’s
Kryptogamen-Flora von Deutschland, 3. Aufl.) 6. Band,
4. Lieferung. Leipzig: Akademische Verlagsgesellschaft;
p. 481–640.
Muñoz J, Cezón K, Hespanhol H, Quandt D. 2015. Grimma.
Flora Briofítica Ibérica. Vol. II. Murcia: Universidad de
Murcia, Sociedad Española de Briología.
Muñoz J, Hespanhol H, Cezón K, Séneca A. 2009. Grimmia
horrida (Grimmiaceae), a new species from the Iberian
Peninsula. The Bryologist. 112:325–328.
Muñoz J, Pando F. 2000. A world synopsis of the genus
Grimmia. Monographs in Systematic Botany from the
Missouri Botanical Garden. 83:1–333.
Nieto-Lugilde M, Werner O, McDaniel SF, Ros RM. 2018.
Environmental variation obscures species diversity in
southern European populations of the moss genus
Ceratodon. Taxon. 67:673–692.
Nieuwkoop J, Arts T. 1995. Additions to the bryophyte flora of
Madeira. Lindbergia. 20:35–39.
O’Shea BJ. 2006. Checklist of the mosses of sub-Saharan Africa
(version 5, 12/06). Tropical Bryology Research Reports.
6:1–253. Available at http://www.nhm.ac.uk/hosted_sites/
bbstbg/resources_lit_africa.html
Ochi H. 1959. A revision of the Bryaceae in Japan and adjacent
regions. Tottori: The Biological Institute, Faculty of Liberal
Arts, Tottori University.
Ochi H. 1970. A revision of the subfamily Bryoideae in
Australia, Tasmania, New Zealand and the adjacent
islands. Journal of the Faculty of Education Tottori
University, Natural Science. 21:7–67.
Ochi H. 1972. A revision of African Bryoidea, Musci (First Part).
Journal of the Faculty of Education Tottori University,
Natural Science. 23:1–126.
Ochi H. 1980. A revision of the Neotropical Bryoideae, Musci
(First part). Journal of the Faculty of Education Tottori
University, Natural Science. 29:49–154.
Ochi H. 1981. A revision of the Neotropical Bryoideae, Musci
(Second part). Journal of the Faculty of Education Tottori
University, Natural Science. 30:21–55.
Ochyra R. 2013. The new generic name Torrentaria, a nomenclatural consequence of the legitimacy of Platyhypnum and
the illegitimacy of Platyhypnidium (Bryophyta). Nova
Hedwigia. 96:205–212.
Ochyra R, Bednarek-Ochyra H. 2011. Lectotypification of
Hypnum boreale (Bryopsida), an early name for a bryalean
species and its taxonomic status. Nova Hedwigia.
93:525–536.
Ochyra R, Bednarek-Ochyra H. 2015. (2384) Proposal to conserve the name Bryum pallescens against Hypnum boreale
(Bryum boreale) (Bryophyta: Bryophyta). Taxon.
64:1053–1054.
Ochyra R, Bednarek-Ochyra H, Plášek V. 2017. Proposal to conserve the name Plagiomnium against Orthomnion and
Orthomniopsis (Bryophyta : Mniaceae). Taxon. 66:1218–1220.
Ochyra R, Gos L. 1992. A new species of Seligeria (Musci,
Seligeriaceae) from Central Europe. Fragmenta Floristica
& Geobotanica. 37:371–378.
Ochyra R, Lewis Smith RI, Bednarek-Ochyra H. 2008. The illustrated moss flora of Antarctica. Cambridge: Cambridge
University Press.
79
Ochyra R, Plášek V, Bednarek-Ochyra H. 2018. Osculatia – an
earlier name for a segregate of Bryum (Bryaceae,
Bryophyta). Acta Musei Silesiae Scientiae Naturales. 67:71–79.
Ochyra R, Stebel A. 2008. Hylocomiadelphus, a new genus of
pleurocarpous mosses. In: Stebel A, Ochyra R, editors.
Bryophytes of the Polish Carpathians. Poznań, Sorus.
p. 303–308.
Ochyra R, Żarnowiec J, Bednarek-Ochyra H. 2003. Census catalogue of Polish mosses. Biodiversity of Poland. 3:1–372.
Olsson S, Buchbender V, Enroth J, Huttunen S, Hedenäs L,
Quandt D. 2009. Evolution of the Neckeraceae
(Bryophyta): resolving the backbone phylogeny.
Systematics and Biodiversity. 7:419–432.
Olsson S, Enroth J, Buchbender V, Hedenäs L, Huttunen S,
Quandt D. 2011. Neckera and Thamnobryum (Neckeraceae,
Bryopsida) – paraphyletic assemblages. Taxon. 60:36–50.
Orgaz JD, Cano MJ, Guerra J. 2013. Taxonomic revision
of Brachytheciastrum (Brachytheciaceae, Bryophyta) from
the Mediterranean Region. Systematic Botany. 38:283–294.
Patiño J, Hedenäs L, Dirkse GM, Ignatov MS, Papp B, Müller F,
González-Mancebo JM, Vanderpoorten A. 2017. Species
delimitation
in
the
recalcitrant
moss
genus
Rhynchostegiella (Brachytheciaceae). Taxon. 66:293–308.
Paton JA, Sheahan MC. 2006. Lophocolea brookwoodiana
(Jungermanniales: Geocalycaceae), a new species in
Britain. Journal of Bryology. 28:163–166.
Pedersen N, Cox CJ, Hedenäs L. 2003. Phylogeny of the moss
family Bryaceae inferred from chloroplast DNA sequences
and morphology. Systematic Botany. 28:471–482.
Pedersen N, Hedenäs L. 2002. Phylogenetic relationships
between Bryum and supposedly closely related genera.
Journal of Bryology. 24:277–289.
Pedersen N, Hedenäs L. 2003. Phylogenetic investigations of a
well supported clade within the acrocarpous moss family
Bryaceae: evidence from seven chloroplast DNA sequences
and morphology. Plant Systematics and Evolution.
240:115–132.
Pedersen N, Hedenäs L. 2005. Taxonomic and nomenclatural
implications of phylogenetic studies of the Bryaceae based
on molecular data and morphology. The Bryologist.
108:123–128.
Pedersen N, Holyoak DT, Newton AE. 2007. Systematics and
morphological evolution within the moss family
Bryaceae: a comparison between parsimony and
Bayesian methods for reconstruction of ancestral character
states. Molecular Phylogenetics and Evolution. 43:891–907.
Perold SM. 1989a. Studies in the genus Riccia (Marchantiales)
from South Africa. 14. R. concava, section Pilifer. Bothalia.
19:161–165.
Perold SM. 1989b. Studies in the genus Riccia (Marchantiales)
from southern Africa. 11. R. montana and R. alboporosa, a
further two new white-scaled species of the group
‘Squamatae’. Bothalia. 19:9–16.
Perold SM. 1997. Studies in the liverwort genus Fossombronia
(Metzgeriales) from southern Africa. 4. A re-examination of
F. crispa, F. leucoxantha and F. tumida. Bothalia. 27:105–115.
Plášek V, Sawicki J, Ochyra R, Szczecińska M, Kulik T. 2015.
New taxonomical arrangement of the traditionally conceived genera Orthotrichum and Ulota. Acta Musei
Silesiae Scientiae Naturales. 64:169–174.
Plášek V, Sawicki J, Trávníčková V, Pasečná M. 2009.
Orthotrichum moravicum (Orthotrichaceae), a new moss
species from the Czech Republic. The Bryologist.
112:329–336.
Podpĕra J. 1954. Conspectus Muscorum Europaeorum.
Prague: Ceskoslovenské Akademie Ved.
80
N. G. HODGETTS ET AL.
Podpěra J. 1973. Bryum generis monographiae prodromus
I. 1. Species Eurasiae septentrionalis. Pars 16. Prague:
Academia, Publishing House of the Czechoslovak
Academy of Sciences.
Porley RD, Edwards S. 2010. Leptodontium proliferum Herzog
(Bryopsida: Pottiaceae), new to Europe. Journal of
Bryology. 32:46–50.
Potemkin AD. 1998. On the origin, evolution and
classification of the genus Scapania (Dum.) Dum.
(Hepaticae). Journal of the Hattori Botanical
Laboratory. 85:33–61.
Potemkin AD. 1999. An analysis of the practical taxonomy of
some critical northern species of Scapania (Scapaniaceae,
Hepaticae). The Bryologist. 102:32–38.
Potemkin AD. 2008. [New liverwort records from Republic of
Komi. 1]. Arctoa 17:198.
Potemkin AD. 2019. The poorly known species Scapania
schljakovii Potemkin (Scapaniaceae, Marchantiophyta), its
gemmae and first record for Bhutan. Journal of Bryology.
41:174–176.
Potemkin AD, Rozantseva EI. 2015. New and little-known for
the Leningrad Region liverworts (Marchantiophyta) from
Hogland Island (Gulf of Finland, Baltic Sea). Novosti sistematiki nizshikh rastenii. 49:369–381.
Price MJ, Ellis LE. 2018. The disentanglement of Andreaea
alpina Hedw. (Andreaeaceae): typifications and nomenclatural changes. Taxon. 67:989–995.
Price MJ, Maier E, Ellis LT. 2019. Dicranum scottianum Turner
(Dicranaceae): synonyms, a type and the correct author.
Journal of Bryology. 41:149–156.
Puche F, Casas C, Brugués M. 2006. Didymodon eckeliae
(Pottiaceae), new to Europe. The Bryologist. 109:239–241.
Puche F, Segarra-Moragues JG. 2013. Riella bialata Trab.
(Riellaceae, Marchantiophyta): a new addition to the
European liverwort flora. Cryptogamie, Bryologie 34:341–
352.
Richards PW. 1936 A collection of bryophytes from the
Azores. Annales Bryologici. 9:131–138.
Rodegher E. 1896. Elenco delle Epatiche della provincia di
Bergamo. Nuovo Giornale Botanico Italiano (n.ser.).
3:423–436.
Rodriguez-Romero A, Hernandez-Ferrer M, Patiño J, Werner
W, González-Mancebo, JM. 2016. Characterization of ten
polymorphic microsatellite loci for the threatened species
Grimmia curviseta Bouman (Grimmiaceae, Musci). Journal
of Bryology. 39:16–32.
Ros RM, Guerra J, Carrion JS, Cano MJ. 1996. A new point of
view on the taxonomy of Pottia starckeana agg. (Musci,
Pottiaceae). Plant Systematics and Evolution. 199:153–165.
Ros RM, Mazimpaka V, Abou-Salama U, Aleffi M, Blockeel TL,
Brugués M, Cano MJ, Cros RM, Dia MG, Dirkse GM, et al.
2007. Hepatics and Anthocerotae of the Mediterranean,
an annotated checklist. Cryptogamie, Bryologie. 28:351–
437.
Ros RM, Mazimpaka V, Abou-Salama U, Aleffi M, Blockeel TL,
Brugués M, Cros RM, Dia MG, Dirkse GM, Draper I, et al.
2013. Mosses of the Mediterranean, an annotated checklist. Cryptogamie, Bryologie. 34:99–283.
Ros RM, Muñoz J, Werner O, Rams S. 2008. New typifications
and synonyms in Tortula sect. Pottia (Ehrh. ex Rchb.) Kindb.
(Pottiaceae, Musci). Taxon. 57:279–288.
Ros RM, Rams S, Holyoak DT, Brugués M, Cros RM. 2007.
Revision of the records of Brachymenium commutatum
(Bryophyta, Bryaceae) in Europe. Nova Hedwigia. 84:241–
247.
Ros RM, Werner O. 2007. The circumscription of the genus
Pottiopsis (Pottiaceae, Bryophyta) based on morphology
and molecular sequence data. Nova Hedwigia, Beiheft.
131:65–79.
Roskov Y, Ower G, Orrell T, Nicolson D, Bailly N, Kirk PM,
Bourgoin T, DeWalt RE, Decock W, Nieukerken E, et al.,
editors. 2019. Species 2000 and ITIS Catalogue of Life,
2019 Annual Checklist. Digital resource at www.
catalogueoflife.org/annual-checklist/2019. Species 2000:
Naturalis, Leiden, the Netherlands. ISSN 2405-884X.
Rycroft DS, Groth H, Heinrichs J. 2004. Reinstatement of
Plagiochila
maderensis
(Jungermanniopsida:
Plagiochilaceae) based on chemical evidence and nrDNA
ITS sequences. Journal of Bryology. 26:37–45.
Saługa M, Ochyra R, Żarnowiec J, Ronikier M. 2018. Do
Antarctic populations represent local or widespread phylogenetic and ecological lineages? Complicated fate of
bipolar moss concepts with Drepanocladus longifolius as
a case study. Organisms Diversity, Evolution. 18:263–278.
Saukel
J.
1985.
Zum
Merkmalsbestand
einiger
mitteleuropäischer
Arten
der
Lebermoosgattung
Lophozia (Dum.) Dum. (Sektion Lophozia). Stapfia.
14:149–185.
Sawicki J, Plášek V, Ochyra R, Szczecińska M, Ślipiko M,
Myszczyński K, Kulik T. 2017. Mitogenomic analyses
support the recent division of the genus Orthotrichum
(Orthotrichaceae, Bryophyta). Scientific Reports. 7:1–10.
Sawicki J, Plášek V, Szczecińska M. 2010. Molecular studies
resolve Nyholmiella (Orthotrichaceae) as a separate
genus. Journal of Systematics and Evolution. 48:183–194.
Sawicki J, Szczecińska M, Bednarek-Ochyra H, Ochyra R. 2015.
Mitochondrial phylogenomics supports splitting the traditionally conceived genus Racomitrium (Bryophyta:
Grimmiaceae). Nova Hedwigia. 100:1–25.
Schill DB, Long DG, Köckinger H. 2008. Taxonomy of Mannia
controversa (Marchantiidae, Aytoniaceae) including a new
subspecies from East Asia. Edinburgh Journal of Botany.
65:35–47.
Schimper WP. 1876. Synopsis Muscorum europaeorum. 2nd
edition. 2 vols. Stuttgartiae: E. Schweizerbart.
Schlesak S, Hedenäs L, Nebel M, Quandt D. 2018. Cleaning a
taxonomic dustbin: placing the European Hypnum
species in a phylogenetic context! Bryophyte Diversity
and Evolution. 40:37–54.
Schlesak S, Hedenäs L, Nebel M, Quandt D. 2019. Addendum
to Hypnum subcomplanatum Hedenäs, Schlesak & Quandt.
Bryophyte Diversity and Evolution. 41:1.
Schljakov RN. 1969. [De positione systematica Lophosiae murmanicae Kaal.]. Novosti sistematiki nizshikh rastenii. 6:241–
245.
Schljakov RN. 1970. [Sectio nova generis Lophozia Dum.
emend. Loeske]. Novosti sistematiki nizshikh rastenii.
7:324–333.
Schljakov RN. 1975. [Notulae systematicae de hepaticis s. str.].
Novosti sistematiki nizshikh rastenii. 12:307–317.
Schljakov RN. 1980. Pechenochnye Mkhi Severa SSSR. 3.
Leningrad: Nauka.
Schljakov RN. 1981. Pechenochnye Mkhi Severa SSSR. 4.
Leningrad: Nauka.
Schljakov RN. 1998. On the Lophozia groenlandica (Nees)
Macoun (Hepaticae). Arctoa. 7:191–196.
Schriebl A. 1991. Experimentelle Studien über die
Laubmoosgattung Polytrichum. Carinthia II. 181/101:461–506.
Schuster RM. 1969. The Hepaticae and Anthocerotae of
North America. II. New York: Columbia University Press.
Schuster RM. 1974. The Hepaticae and Anthocerotae of North
America. III. New York: Columbia University Press.
Schuster RM. 1988. The hepaticae of South Greenland.
Beihefte zur Nova Hedwigia. 92:1–255.
JOURNAL OF BRYOLOGY
Schuster RM. 1992a. Studies on Marchantiales. I–III. Journal of
the Hattori Botanical Laboratory. 71:267–287.
Schuster RM. 1992b. The Hepaticae and Anthocerotae of
North America. VI. New York: Columbia University Press.
Schwägrichen CF. 1816. Species Muscorum Frondosorum,
Supplementum Primum. Leipzig: Barth.
Schwägrichen CF. 1827. Species Muscorum Frondosorum,
Supplementum Tertium, vol. 1, sect. 1. Leipzig: Barth.
Segarra-Moragues JG, Puche F, Sabovljević M, Infante M,
Heras P. 2014. Taxonomic revision of Riella subgenus
Trabutiella (Riellaceae, Sphaerocarpales). Phytotaxa.
159:131–174.
Séneca A, Söderström L. 2009. Sphagnophyta of Europe and
Macaronesia: a checklist with distribution data. Journal of
Bryology. 31:243–254.
Sérgio C. 1985. Notulae Bryoflorae Lusitanicae I. 5 - Notas
acerca do género Fossombronia Raddi em Portugal.
Portugaliae Acta Biologica. Série B. Sistemática, Ecologia,
Biogeografia e Paleontologia. 14:188–194.
Sérgio C. 2003. Notulae Bryoflorae Lusitanicae IX. 1. Acerca sa
precença de Fossombronia leucoxantha Lehm. e F. crispa
Ness [sic!] em Portugal. Portugaliae Acta Biologica. Série
B. Sistemática, Ecologia, Biogeografia e Paleontologia.
21:279–282.
Sérgio C. 2006. A review of the Gymnostomum calcareum
Nees & Hornsch. complex (Bryopsida: Pottiaceae) in
southern Europe and the Macaronesian Islands, including
G calcareum var. atlanticum var. nov. Journal of Bryology.
28:38–45.
Sérgio C, Garcia, C. 2009. Noteworthy range extensions of two
Aneura (Jungermanniopsida, Metzgeriales) species new to
the Iberian Peninsula: Aneura maxima (Schiffn.) Steph. and
A. pseudopinguis (Herzog) Pócs. Cryptogamie, Bryologie.
30:207–215.
Sérgio C, Sim-Sim M. 2012. Andreaea flexuosa R.Brown bis
subsp. luisieri Sérgio & Sim-Sim (Andreaeaceae), a new
taxon from Madeira Island. Cryptogamie, Bryologie.
33:271–277.
Sérgio C, Sim-Sim M, Luis L. 2010. Atrichum androgynum
(Müll.Hal.) A.Jaeger in Portugal, Azores and Madeira, new
to the European bryoflora. Cryptogamie, Bryologie.
31:281–288.
Shaw AJ. 2014. The impact of molecular data on our understanding of the bryophyte flora of North America. Flora
of North America Editorial Committee, editors. Flora of
North America. Vol. 28. Bryophyta, Part 2. New York:
Oxford University Press; p. 3–13.
Shaw AJ, Flatberg KI, Szövényi P, Ricca M, Johnson MG,
Stenøien HK, Shaw B. 2012. Systematics of the Sphagnum
fimbriatum complex: phylogenetic relationships, morphological variation, and allopolyploidy. Systematic Botany.
37:15–30.
Shaw B, Crandall-Stotler B, Váňa J, Stotler RE, von Konrat M,
Engel JJ, Davis CE, Long DG, Sova P, Shaw AJ. 2015.
Phylogenetic relationships and morphological evolution
in a major clade of leafy liverworts (phylum
Marchantiophyta, order Jungermanniales): suborder
Jungermanniineae. Systematic Botany. 40:27–45.
Simmel J, Poschlod P. 2017. Beiträge zur Bestimmung und
Öckologie der mitteleuropäischen Leucobryum-Taxa.
Herzogia. 30:397–411.
Sim-Sim M. 1999. The genus Frullania Raddi (Hepaticae) in
Portugal and Madeira. Cryptogamie, Bryologie 20:83–
144.
Sim-Sim M, Afonina OM, Almeida T, Désamoré A, Laenen
B, Garcia CA, González-Mancebo JM, Stech M. 2017.
Integrative
taxonomy
reveals
too
extensive
81
lumping and a new species in the moss genus
Amphidium (Bryophyta). Systematics and Biodiversity.
15:451–463.
Skrzypczak R. 2001. Découverte de Riella parisii Gottsche dans
le Var (espèce nouvelle pour la France) et d’une deuxième
station de Riella helicophylla (Bory & Mont.) Mont. dans
l’Hérault. Bulletin de la Société Botanique du CentreOuest, nouvelle série. 32:277–290.
Smith AJE. 1990. The liverworts of Britain and Ireland.
Cambridge: Cambridge University Press.
Smith AJE. 2004. The moss flora of Britain and Ireland. 2nd ed.
Cambridge: Cambridge University Press.
Söderström L, De Roo RT, Hedderson TAJ. 2010. Taxonomic
novelties resulting from recent reclassification of the
Lophoziaceae/Scapaniaceae clade. Phytotaxa. 3:47–53.
Söderström L, Hagborg A, von Konrat M. 2019. ELPT: Early
Land Plants Today (version Jan 2019). In: Roskov Y, Ower
G, Orrell T, Nicolson D, Bailly N, Kirk PM, Bourgoin T,
DeWalt RE, Decock W, Nieukerken E, et al., editors.
Species 2000 and ITIS Catalogue of Life, 2019 Annual
Checklist. Digital resource at www.catalogueoflife.org/
annual-checklist/2019. Species 2000: Naturalis, Leiden,
the Netherlands. ISSN 2405-884X.
Söderström L, Hagborg A, von Konrat M, Bartholomew-Began
S, Bell D, Briscoe L, Brown E, Cargill DC, Costa DP, CrandallStotler BJ, et al. 2016. World checklist of hornworts and
liverworts. PhytoKeys. 59:1–828.
Söderström L, Urmi E, Váňa J. 2002. Distribution of Hepaticae
and Anthocerotae in Europe and Macaronesia. Lindbergia.
27:3–47.
Söderström L, Urmi E, Váňa J. 2007. The distribution of hepaticae and anthocerotae in Europe and Macaronesia –
Update 1-427. Cryptogamie, Bryologie. 28:299–350.
Söderström L, Váňa J, Crandall-Stotler B, Hentschel J, Hagborg
A, von Konrat MJ. 2015. Notes on Early Land Plants Today.
68. Miscellaneous notes on Marchantiophyta. Phytotaxa.
202:69–72.
Söderström L, Váňa J, Hagborg A, von Konrat M. 2013. Notes
on Early Land Plants Today. 35. Notes on Lophoziaceae
(Marchantiophyta). Phytotaxa. 97:27–35.
Sofronova EV, Abakarova AS, Afonina OV, Akatova TV,
Baisheva EZ, Baj X-L, Bezgodov AG, Bochkin VD,
Borovichev EA, Chernadjeva IV, et al. 2013. New bryophyte
records. 2 — Novye briologičeskie nahdki. 2. Arctoa
22:239–262.
Sotiaux A, Enroth J, Olsson S, Quandt D, Vanderpoorten A.
2009. When morphology and molecules tell us different
stories: a case-in-point with Leptodon corsicus, a new and
unique endemic moss species from Corsica. Journal of
Bryology. 31:186–196.
Spence JR. 2005. New genera and combinations in Bryaceae
(Bryales, Musci) for North America. Phytologia. 87:15–28.
Spence JR. 2014. Bryaceae. In: Flora of North America Editorial
Committee, editors. Flora of North America. Vol. 28.
Bryophyta, Part 2. New York: Oxford University Press.
Sprengel KPJ. 1807. Mantissa Prima Florae Halensis. C.
Kumelium: Halae [Halle].
Spruce R. 1876a. On Anomoclada, a new genus of
Hepaticae, and on its allied genera, Odontoschisma
and Adelanthus (cont.). Journal of Botany, British and
Foreign. 14:193–203.
Spruce R. 1876b. On Anomoclada, a new genus of hepaticae,
and on its allied genera, Odontoschisma and Adelanthus
(concluded). Journal of Botany, British and Foreign.
14:230–235.
Stech M. 1999. Dichodontium palustre (Dicks.) Stech comb.
nov., a new name for Dicranella palustris (Dicks.) Crundw.
82
N. G. HODGETTS ET AL.
ex Warb. (Dicranaceae, Bryopsida). Nova Hedwigia. 69:237–
240.
Stech M, McDaniel SF, Hernández-Maqueda R, Ros RM,
Werner O, Muňoz J, Quandt D. 2012. Phylogeny of haplolepidous mosses – challenges and perspectives. Journal of
Bryology. 34:173–186.
Stech M, Sim-Sim M, Esquível MG, Fontinha S, Tangney R,
Lobo C, Gabriel R, Quandt D. 2008. Explaining the ‘anomalous’ distribution of Echinodium (Bryopsida: Echinodiaceae):
independent evolution in Macaronesia and Australasia.
Organisms, Diversity and Evolution. 8:282–292.
Ştefănuţ S. 2008. The hornwort and liverwort atlas of
Romania. Bucharest: Ars Docendi.
Stephani F. 1903. Species hepaticarum 2. Bulletin de l’Herbier
Boissier (sér. 2) 3:326–341.
Stephani F. 1905. Species hepaticarum 3. Bulletin de l’Herbier
Boissier (sér. 2). 5:1129–1144.
Stephani F. 1907. Species hepaticarum 3. Bulletin de l’Herbier
Boissier (sér. 2). 7:477–492.
Stotler RE, Bray JR jr, Cargill DC, Krayesky D, Crandall-Stotler
BJ. 2003. Typifications in the genus Fossombronia
(Marchantiophyta). The Bryologist. 106:130–142.
Stotler RE, Crandall-Stotler BJ. 2017. A synopsis of the liverwort flora of North America north of Mexico. Annales of
the Missouri Botanical Garden. 102:574–709.
Streiff A. 2006. Phylogenetic study of Grimmia
(Grimmiaceae) based on plastid DNA sequences (trnLtrnF and rps4) and on morphological characters. The
Bryologist. 109:224–235.
Sun Y, He X. 2019. On the identity of Herbertus aduncus
(Dicks.) Gray, a taxonomic update. Acta Bryolichenologica
Asiatica. 8:145–161.
Szweykowski J, Buczkowska K, Odrzykoski IJ. 2005.
Conocephalum
salebrosum
(Marchantiopsida,
Conocephalaceae) – a new holarctic liverwort species.
Plant Systematics and Evolution. 253:133–158.
Tassi A. 1901. Epatiche coltivate nell’Orto botanico di Siena.
Bullettino del laboratorio ed orto botanico della
R. università di Siena. 4:17–18.
Tubanova DY, Goryunov DV, Ignatova EA, Ignatov MS. 2010. On
the taxonomy of Dicranum acutifolium and D. fuscescens complexes (Dicranaceae, Bryophyta) in Russia. Arctoa. 19:151–164.
Tubanova DY, Ignatova EA. 2011. A new species of Dicranum
(Dicranceae, Bryophyta) from Asiatic Russia. Arctoa.
20:183–190.
Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL,
Herendeen PS, Knapp S, Kusber W-H, Li D-Z, Marhold K, et al.
2018. International Code of Nomenclature for algae, fungi,
and plants (Shenzhen Code). Regnum Vegetabile. 159:1–254.
Ulvinen T, Kypärä T. 2016. Nipponinlähdesammal (Philonotis
yezoana) Suomen Lapissa, Bryobrotherella. 19:71–76.
Unger FJAN, Klotschy CGT 1865. Die Insel Cypern. Wien:
Wilhelm Braun Müller.
Urmi E. 2017. Betreffend Diplophyllum obtusatum
(R.M.Schust.) R.M.Schust. in der Schweiz. Meylania. 60:5–7.
Váňa J. 1986. Ochyraea tatrensis gen. et spec. nov., a remarkable pleurocarpous moss from Czechoslovakia. Journal of
Bryology. 14:261–267.
Váňa J, Bechteler J, Schäfer-Verwimp A, Renner MAM, Heinrichs
J. 2017. Integrative taxonomy substantiates the presence
of three Radula species in Austria: Radula complanata,
R. lindenbergiana, and R. visianica (Porellales,
Jungermanniopsida). Cryptogamie, Bryologie 38:125–135.
Váňa J, Engel JJ. 2013. The liverworts and hornworts of the
Tristan da Cunha group of islands in the south Atlantic
Ocean. Memoirs of the New York Botanical Garden.
105:1–138.
Váňa J, Söderström L, Hagborg A, von Konrat M. 2013. Notes
on Early Land Plants Today. 40. Notes on Cephaloziellaceae
(Marchantiophyta). Phytotaxa 112:1–6.
Vanderpoorten A. 2004. A simple taxonomic treatment for a
complicated
evolutionary
story:
the
genus
Hygroamblystegium
(Hypnales,
Amblystegiaceae).
Monographs in Systematic Botany from the Missouri
Botanical Garden. 98:320–327.
Vanderpoorten A, Boles S, Shaw AJ. 2003. Patterns of
molecular and morphological variation in Leucobryum
albidum, L. glaucum, and L. juniperoideum (Bryopsida).
Systematic Botany. 28:651–656.
Vanderpoorten A, Hedenäs L. 2009. New combinations in the
Amblystegiaceae. Journal of Bryology. 31:129–132.
Vanderpoorten A, Hedenäs L, Cox CJ, Shaw AJ. 2002.
Circumscription, classification, and taxonomy of
Amblystegiaceae (Bryopsida) inferred from nuclear and
chloroplast DNA sequence data and morphology. Taxon.
51:115–122.
Vanderpoorten A, Patiño J, Dirkse G, Blockeel T, Hedenäs L.
2015. Early divergence of an Azorean endemic species in
the moss genus Rhynchostegiella (Brachytheciaceae).
Phytotaxa. 210:60–69.
Vigalondo B, Garilleti R, Vanderpoorten A, Patiño J, Draper I,
Calleja JA, Mazimpaka, V, Lara F. 2019. Do mosses really
exhibit so large distribution ranges? Insights from the integrative taxonomic study of the Lewinskya affinis complex
(Orthotrichaceae, Bryopsida). Molecular Phylogenetics
and Evolution. 140:106598.
Vigalondo B, Patiño J, Draper I, Mazimpaka V, Shevock JR,
Losada-Lima A, González-Mancebo JM, Garilleti R, Lara F.
2019. The long journey of Orthotrichum shevockii
(Orthotrichaceae, Bryopsida): from California to
Macaronesia. PLoS One. 14(2):e0211017.
Villarreal JC, Forrest LL, Cooper ED, Long DG. 2014.
Phylogenetic affinities and conservation status of
Telaranea murphyae Paton in Britain. Journal of Bryology.
36:191–199.
Vilnet AA, Borovichev EA, Bakalin VA. 2014. Frullania subarctica - a new species of the Frullania tamarisci complex
(Frullaniaceae, Marchantiophyta). Phytotaxa. 173:61–72.
Vilnet AA, Konstantinova NA, Troitsky AV. 2010. Molecular
insight on phylogeny and systematics of the
Lophoziaceae, Scapaniaceae, Gymnomitriaceae and
Jungermanniaceae. Arctoa. 19:31–50.
Vilnet AA, Milyutina IA, Troitsky AV, Konstantinova NA. 2005.
[On phylogeny and systematics of genus Lophozia
(Dumort.) Dumort. s.str. based on trnL-trnF sequences of
chloroplast DNA and ITS 1-2 sequences of nuclear DNA].
In: Vilnet AA, Milyutina IA, Troitsky AV, Konstantinova NA,
editors. Actual problems in Bryology: proceedings of the
international meeting devoted to the 90th anniversary of
A. L. Abramova (Saint Petersburg, November 22–25, 2005).
St. Petersburg: Komarov Botanical Institute; p. 37–43.
Vitt DH, Varigo C. 1997. Orthotrichum elegans, a taxon worthy
of species rank. Journal of the Hattori Botanical Laboratory.
82:329–335.
Wang CY, Zhao JC. 2009. Phylogeny of Ptychostomum
(Bryaceae, Musci) inferred from sequences of nuclear ribosomal DNA internal transcribed spacer (ITS) and chloroplast
rps4. Journal of Systematics and Evolution. 47:311–320.
Werner O, Köckinger H, Magdy M, Ros RM. 2014. On the systematic position of Tortella arctica and Trichostomum arcticum (Bryophyta, Pottiaceae). Nova Hedwigia. 98:273–293.
Werner O, Patiño J, González–Mancebo JM, Gabriel R, Ros RM.
2009. The taxonomic status and the geographical relationships of the Macaronesian endemic moss Fissidens luisieri
JOURNAL OF BRYOLOGY
83
(Fissidentaceae) based on DNA sequence data. The
Bryologist. 112:315–324.
Werner O, Rams S, Kučera J, Larraín J, Afonina OM, Pisa S, Ros
RM. 2013. New data on the moss genus Hymenoloma, with
special reference to H. mulahaceni. Cryptogamie,
Bryologie. 34:13–30.
Werner O, Rodriguez-Atienza A, Nieto-Lugilde M, Ros RM.
2015. New data in the systematics of the European-Asian
species of Leucodon (Leucodontaceae, Bryophyta). Nova
Hedwigia. 100:333–354.
Werner O, Ros RM, Goffinet B. 2007. A reconsideration of
the systematic position of Goniomitrium (Funariaceae)
based on chloroplast sequence markers. The Bryologist.
110:108–114.
Werner O, Ros RM, Grundmann M. 2005. Molecular phylogeny
of Trichostomoideae (Pottiaceae, Bryophyta) based on
nrITS sequence data. Taxon. 54:361–368.
Withering W. 1776. A botanical arrangement of all the vegetables naturally growing in Great Britain. Vol. II.
Birmingham: M. Swinney; p. 385–838.
Wynns JT, Schröck C. 2018. Range extensions for the rare
moss Plagiothecium handelii, and its transfer to the resurrected genus Ortholimnobium. Lindbergia. 41:1–7.
Zander RH. 2013. A framework for post-phylogenetic systematics. St. Louis: Zetetic Publications. CreateSpace
Independent Publishing, Amazon
Zander RH. 2019. Macroevolutionary versus molecular
analysis: systematics of the Didymodon segregates
Aithobryum, Exobryum and Fuscobryum (Pottiaceae).
Hattoria. 10:1–38.
Zander RH, Hedderson TA. 2016. Re-evaluation of
Hymenostylium xanthocarpum (Hook.) Brid., and Ardeuma
R.H.Zander & Hedd., a new name for all other species of
Hymenostylium (Pottiaceae, Bryophyta). Journal of
Bryology. 38:295–301.
Zinovjeva LA. 1973. K flore pechenochnich mchov poljarnogo i severnogo Urala [On the flora of hepatics of
Polar and North Urals]. Ucenye Zapiski. Permskij
Gosudarstvennyj Universitet imeni A. M. Gor’kogo.
263:14–37.
Zolotov VI. 2007a. On systematics and distribution of some
species of Bryum (Bryaceae, Bryophyta) in Russia. Arctoa.
15:155–162.
Zolotov VI 2007b. Genus 1. Bryum Hedw. – Brium, p. 1–67.
http://arctoa.ru/Flora/taxonomy-ru/Bryum1.htm. [Accessed
2010 May 25].
Index
A. lindenbergianus 9
Aithobryum = Didymodon 37
A. bistratosum = Didymodon bistratosus 37
Alleniella 64
A. besseri 64
A. complanata 64
Aloina 36
A. aloides 36
A. ambigua 36
A. bifrons 36
A. brevirostris 36
A. humilis 36
A. obliquifolia 36
A. rigida 36
Alophosia 29
A. azorica 29
Amblyodon 48
A. dealbatus 48
AMBLYSTEGIACEAE 2, 8, 56, 57
Amblystegioideae 56
Amblystegium 56
A. confervoides = Serpoleskea confervoides 58
A. radicale = Pseudocampylium radicale 58
A. serpens 56
var. salinum = Amblystegium serpens 56
A. subtile = Pseudoamblystegium subtile 58
A. varium = Hygroamblystegium varium 57
AMPHIDIACEAE 7, 33
Amphidium 33
A. curvipes 33, 68
A. lapponicum 33
A. mougeotii 33
[A. tortuosum] 68
A. tortuosum auct. eur. = Amphidium curvipes 33
Anacamptodon 56, 57
A. sauteri = Microhypnum sauteri 57
A. splachnoides 56
Anacolia 47
A. laevisphaera = Bartramia laevisphaera 47
A. menziesii 47
A. webbii 47
Anastrepta 9
This index enumerates all names mentioned in the text,
accepted taxa as well as synonyms. Taxa in bold are accepted
taxa occurring in Europe or northern Macaronesia and taxa in
square brackets are taxa only occurring outside the area.
Abietinella 59
A. abietina 59
var. abietina 59
var. hystricosa 59
Acanthocoleus 22
A. aberrans 22
[var. aberrans] 22
var. laevis 22
Acaulon 36
A. casasianum 36
A. dertosense 36
A. fontiquerianum 36
[A. longifolium] 69
A. mediterraneum 36
A. muticum 36
A. piligerum 36
A. triquetrum 36
Achrophyllum 55
A. dentatum 4, 55
ACROBOLBACEAE 6, 15
Acrobolboideae 15
Acrobolbus 16
[A. anisodontus] 66
A. azoricus 16, 66
[A. ciliatus] 66
[A. laxus] 66
A. madeirensis 16, 66
A. wilsonii 16, 66
[var. andinus] 16
var. wilsonii 16
[Acrolejeunea emergens] 68
ADELANTHACEAE 6, 9
Adelanthoideae 9
Adelanthus 9
A. section Lindenbergiani 9
A. decipiens = Pseudomarsupidium decipiens 9
84
N. G. HODGETTS ET AL.
A. orcadensis 9
ANASTROPHYLLACEAE 6, 9
Anastrophyllum 9, 10
A. alpinum 9
A. assimile 9
var. nardioides = Anastrophyllum assimile 9
A. cavifolium = Orthocaulis cavifolius 10
A. donnianum 9
A. hellerianum = Crossocalyx hellerianus 9
A. joergensenii 9
A. michauxii 9
A. minutum = Sphenolobus minutus 10
var. weberi = Sphenolobus minutus 10
A. saxicola = Sphenolobus saxicola 10
A. sphenoloboides = Schizophyllopsis sphenoloboides 10
Andoa 65
A. berthelotiana 65
Andreaea 29
A. section Andreaea 29
A. section Chasmocalyx 29
A. section Nerviae 29
A. alpestris 29
A. alpina 29
A. alpina auct. mult. = Andreaea hookeri 29
A. blyttii 29
A. crassinervia 29
A. flexuosa 29
[subsp. flexuosa] 29
subsp. luisieri 29
A. frigida 29
A. hartmanii = Andreaea alpina 29
A. heinemannii 29
subsp. crassifolia 29
subsp. heinemannii 29
A. hookeri 29
A. megistospora 29
A. mutabilis 29
A. nivalis 29
A. obovata = Andreaea alpina 29
A. rothii 29
subsp. falcata 29
subsp. rothii 29
A. rupestris 29
var. alpestris = Andreaea alpestris 29
var. papillosa 29
var. rupestris 29
A. sinuosa 29
ANDREAEACEAE 7, 29
ANDREAEALES 7, 29
Andreaeidae 7
Andreaeopsida 7, 29
Aneura 23
A. latissima 23
A. maxima 23
A. mirabilis 23
A. pinguis 23
var. angustior = Aneura pinguis 23
var. denticulata = Aneura pinguis 23
var. fuscovirens = Aneura pinguis 23
A. pseudopinguis = A. latissima 23
ANEURACEAE 6, 23
Anisothecium palustre = Diobelonella palustris 33
Anoectangium 40
A. aestivum 40
A. angustifolium 40
A. handelii 40
A. hornschuchianum = Molendoa hornschuchiana 42
A. sendtnerianum = Molendoa hornschuchiana 42
A. schliephackii = Molendoa schliephackei 42
A. taeniatifolium = Molendoa taeniatifolia 42
A. warburgii = Molendoa warburgii 42
Anomobryum 3, 48
A. apiculatum 4, 5, 48
A. bavaricum = Bryum bavaricum 48
A. concinnatum 48
A. julaceum 48
A. lusitanicum 48
A. notarisii 5, 48
Anomodon 66
A. attenuatus = Pseudanomodon attenuatus 64, 66
A. longifolius 66
A. rostratus = Claopodium rostratum 58
A. rugelii 66
A. tristis 66
A. viticulosus 66
ANOMODONTACEAE 8, 66
Anomodontella = Anomodon 66
A. longifolia = Anomodon longifolius 66
Anomodontopsis = Anomodon 66
A. rugelii = Anomodon rugelii 66
Anthelia 16
A. julacea 16
A. juratzkana 16
ANTHELIACEAE 6, 16
Anthoceros 8
A. agrestis 8
A. caucasicus 8
A. neesii 8
A. punctatus 8
subsp. agrestis = Anthoceros agrestis 8
var. douinii = Anthoceros agrestis 8
ANTHOCEROTACEAE 6, 8
ANTHOCEROTALES 6, 8
Anthocerotidae 6
ANTHOCEROTOPHYTA 2, 6, 8
ANTHOCEROTOPSIDA 6, 8
Antitrichia 64
A. californica 64
A. curtipendula 64
ANTITRICHIACEAE 8, 64
Aongstroemia 33
A. longipes 33
AONGSTROEMIACEAE 7, 33
Aphanolejeunea = Cololejeunea 22
A. azorica = Cololejeunea azorica 22
A. madeirensis = Cololejeunea madeirensis 22
A. microscopica = Cololejeunea microscopica 22
A. sintenisii = Cololejeunea sintenisii 22
Aphanorrhegma = Physcomitrium 31
A. patens = Physcomitrium patens 31
Aplodon 48
A. wormskioldii 48
Apomarsupella = Gymnomitrion 16
A. revoluta = Gymnomitrion revolutum 16
Apometzgeria = Metzgeria 24
A. pubescens = Metzgeria pubescens 24
Apopellia 24
A. endiviifolia 24
Aquilonium 63
A. plicatulum 63
ARCHIDIACEAE 7, 32
Archidium 32
A. alternifolium 32
Arctoa 35
JOURNAL OF BRYOLOGY
A. anderssonii 35
A. fulvella 35
A. hyperborea 35
Ardeuma = Hymenostylium 42
A. annotinum = Hymenostylium recurvirostrum var.
insigne 42
A. insigne = Hymenostylium recurvirostrum var. insigne 42
A. recurvirostrum = Hymenostylium recurvirostrum var. recurvirostrum 42
Arnellia 16
A. fennica 16
ARNELLIACEAE 6, 16
Arvernella 56
A. microclada 56
Aschisma 40
A. carniolicum 40
A. cuynetii 40
Asterella = [Fimbraria] 25
A. subgenus Asterella 25
A. subgenus Phragmoblepharis 25
A. subgenus Saccatae 25
A. section Brachyblepharis 25
A. africana 25
[A. blumeana] 66
[A. elegans] 66
A. gracilis = Mannia gracilis 25
A. lindenbergiana 25
[A. marginata] 66
[A. pringlei] 66
A. saccata 25
[A. tenella] 66
[A. wallachiana] 66
Athalamia auct. eur. = Clevea 25
A. hyalina = Clevea hyalina 25
var. suecica = Athalamia hyalina 25
A. spathysii = Clevea spathysii 25
Atractylocarpus 32
A. alpinus 32
A. subporodictyon 32
Atrichum 29
[A. acuminatum] 69
A. androgynum 29
A. angustatum 29
A. crispum 4, 29
A. flavisetum 29
A. tenellum 29
A. undulatum 29
AULACOMNIACEAE 8, 55
AULACOMNIALES 8, 55
Aulacomnium 55
[A. acuminatum] 69
A. androgynum 55
A. palustre 55
A. turgidum 55
AYTONIACEAE 7, 25
Barbilophozia 9
B. subgenus Barbilophozia 9
B. subgenus Sudeticae 9
B. atlantica = Orthocaulis atlanticus 10
B. attenuata = Neoorthocaulis attenuatus 9
B. barbata 9
B. binsteadii = Neoorthocaulis binsteadii 10
B. floerkei = Neoorthocaulis floerkei 10
B. hatcheri 9
B. hyperborea = Neoorthocaulis hyperboreus 10
B. kunzeana = Schljakovia kunzeana 10
B. lycopodioides 9
B. quadriloba = Schljakovianthus quadrilobus 10
B. rubescens 9
B. sudetica 9
Barbula 36
B. amplexifolia = Hydrogonium amplexifolium 41
B. bicolor = Gymnobarbula bicolor 41
B. bolleana = Hydrogonium bolleanum 41
B. consanguinea = Hydrogonium consanguineum 41
B. convoluta = Streblotrichum convolutum 40
var. sardoa = Streblotrichum convolutum var.
commutatum 40
B. crocea = Hydrogonium croceum 41
B. enderesii = Streblotrichum enderesii 40
[B. indica] 41, 68
B. unguiculata 36
Barbuloideae = Pottioideae 36
Bartramia 47
B. section Bartramia 47
B. section Pyridium 47
B. section Strictidium 47
B. aprica 47, 68
B. breviseta 47
B. halleriana 47
B. ithyphylla 47
B. laevisphaera 47
B. pomiformis 47
B. rosamrosiae = Bartramia aprica 47
[B. stricta] 68
B. stricta auct. eur.= Bartramia aprica 47
B. subulata 47
BARTRAMIACEAE 7, 47
BARTRAMIALES 7, 47
Bartramioideae 47
Bazzania 19
B. azorica 19
[B. denudata] 66
B. flaccida 19
B. pearsonii 19
B. tricrenata 19
[var. fulfordiae] 19
var. tricrenata 19
B. trilobata 19
var. depauperata 19
var. trilobata 19
Bazzanioideae 19
Biantheridion 9
B. undulifolium 9
Blasia 25
B. pusilla 25
BLASIACEAE 7, 25
BLASIALES 7, 25
Blasiidae 7
Blepharostoma 19
[B. arachnoideum] 66
B. trichophyllum 19
subsp. brevirete 19, 66
subsp. trichophyllum 19
var. brevirete = Blepharostoma trichophyllum
subsp. brevirete 19
BLEPHAROSTOMATACEAE 6, 19
Blindia 43
B. acuta 43
B. caespiticia 43
Blindiadelphus 43
B. campylopodus 43
B. diversifolius 43
B. polaris 43
85
86
N. G. HODGETTS ET AL.
B. recurvatus 43
B. subimmersus 43
Brachiolejeuneeae (trib.) 22
Brachiolejeuneinae (subtrib.) 22
Brachydontium 43
B. trichodes 44
Brachymenium 3, 48, 49, 50, 60
[B. acuminatum] 69
[B. commutatum] 68, 69
[B. exile] 69
B. notarisii = Anomobryum notarisii 48
B. paradoxum 48
B. philonotula = Bryum kikuyense 49
BRACHYTHECIACEAE 8, 59
Brachytheciastrum 60
[B. bellicum] 69
B. collinum 60, 68
B. dieckei 60
[B. fendleri] 60, 68
B. fendleri auct. eur. = Brachytheciastrum collinum 60
B. olympicum 60
B. salicinum 60
B. trachypodium 60
[B. umbilicatum] 69
B. velutinum 60
var. vagans = Brachytheciastrum velutinum 61
var. salicinum = Brachytheciastrum salicinum 60
B. vanekii = Brachytheciastrum velutinum 61
Brachythecioideae 60
Brachythecium 61
B. albicans 61
B. buchananii 61
B. campestre 61
B. capillaceum 61
B. cirrosum 61
B. coruscum = Brachythecium tauriscorum 61
B. erythrorrhizon 61
subsp. asiaticum = Brachythecium erythrorrhizon 61
var. thedenii = Brachythecium erythrorrhizon 61
B. funkii 61
B. geheebii 61
B. glareosum 61
B. japygum 61
B. laetum 61
B. mildeanum 61
var. udum = Brachythecium udum 61
B. novae-angliae 61
B. percurrens = Hedenasiastrum percurrens 60
B. rotaeanum = Brachythecium capillaceum 61
B. rivulare 61
B. rutabulum 61
var. atlanticum 61
var. rutabulum 61
B. salebrosum 61
B. scabridum = Brachythecium novae-angliae 61
B. tauriscorum 61
B. tenuicaule 61
B. tommasinii 61
B. turgidum 61
B. udum 61
Braunia 46, 47
B. alopecura 47
B. imberbis 47
Breidleria = Stereodon 62
B. pratensis = Stereodon pratensis 63
Breutelia 47
B. azorica 47
B. chrysocoma 47
Breutelioideae 47
Brotherella 62
[B. henonii] 62, 68
B. henonii auct. eur. = Brotherella lorentziana 62
B. lorentziana 62, 68
Bruchia 36
B. flexuosa 36
B. vogesiaca 36
BRUCHIACEAE 7, 36
BRYALES 8, 48
BRYACEAE 2, 3, 8, 48, 50
Bryhnia = Brachythecium 61
B. novae-angliae = Brachythecium novae-angliae 61
B. scabrida = Brachythecium novae-angliae 61
Bryidae 7
Bryobrittonia 30
B. longipes 30
Bryoerythrophyllum 37
B. alpigenum 37
B. caledonicum 37
B. campylocarpum 37
B. duellii 37
B. ferruginascens 37
B. inaequalifolium 37
B. recurvirostrum 37
B. rubrum 37
Bryophyta 7, 27
Bryophytina 7
Bryopsida 7, 30
BRYOXIPHIACEAE 7, 32
Bryoxiphium 32
B. madeirense 32
B. norvegicum 32
Bryum 3, 48
B. algovicum = Ptychostomum compactum 50
B. algovicum var. rutheanum = Ptychostomum algovicum var.
rutheanum 50
B. alpinum = Imbribryum alpinum 49
[B. anomodon] 69
B. apiculatum = Anomobryum apiculatum 5, 48
B. archangelicum = Ptychostomum inclinatum 50
B. arcticum = Ptychostomum arcticum 50
B. argenteum 48
[B. atrovirens] 69
B. austriacum = Ptychostomum austriacum 6, 50
B. barnesii = Bryum dichotomum 48, 49
B. bavaricum 48
B. blindii 48
B. boreale = Ptychostomum pallescens 51
B. bornholmense = Ptychostomum bornholmense 50
B. caespiticium = Ptychostomum imbricatulum 3, 50
B. calophyllum = Ptychostomum calophyllum 50
B. canariense 48
B. capillare = Ptychostomum capillare 50
B. caucasicum 48
B. cellulare = Ptychostomum cellulare 6, 50
B. compactum = Ptychostomum compactum 50
B. creberrimum = Ptychostomum creberrimum 50
B. cryophilum = Ptychostomum cryophilum 50
B. cyclophyllum = Ptychostomum cyclophyllum 50
B. demaretianum 48
B. dichotomum 3, 48, 49
B. dixonii 49
B. donianum = Ptychostomum donianum 50
B. dyffrynense 49
B. elegans = Ptychostomum elegans 6, 50
JOURNAL OF BRYOLOGY
B. funkii = Ptychostomum funkii 50
B. gemmiferum 49
B. gemmilucens 49
B. gemmiparum 49
B. inclinatum = Distichium inclinatum 50
B. intermedium = Ptychostomum intermedium 50
B. kikuyense 49, 68
B. klinggraeffii 49
B. knowltonii = Ptychostomum knowltonii 50
B. kunzei = Ptychostomum kunzei 50, 51
B. longisetum = Ptychostomum longisetum 51
B. marratii 49
B. mildeanum = Imbribryum mildeanum 49
B. miniatum = Imbribryum miniatum 49
B. minii = Ptychostomum minii 6, 51
B. moravicum = Ptychostomum moravicum 51
B. muehlenbeckii = Imbribryum muehlenbeckii 49
B. neodamense = Ptychostomum pseudotriquetrum 51
B. notarisii = Anomobryum notarisii 5
B. oblongum 49
B. pallens = Ptychostomum pallens 51
[subsp. acutiusculum = Ptychostomum pallens] 51
subsp. prosboreum = Ptychostomum pallens 51
subsp. sibiricum = Ptychostomum pallens 51
f. gracile = Ptychostomum pallens 51
B. pallescens = Ptychostomum pallescens 51
[B. philonotula Müll.Hal.] 68
B. philonotula Hampe = Bryum kikuyense 49
[B. philonotulum] 49, 68
B. philonotulum auct. eur. = Bryum kikuyense 49
B. pseudotriquetrum = Ptychostomum pseudotriquetrum 51
var. bimum = Ptychostomum pseudotriquetrum var.
bimum 51
B. radiculosum 49
B. reyeri 49
B. riparium 49
B. rubens = Ptychostomum rubens 51
B. ruderale 49
B. salinum = Ptychostomum salinum 51
B. sauteri 49
B. schleicheri = Ptychostomum schleicheri 6, 51
var. latifolium = Ptychostomum schleicheri var.
latifolium 6, 51
B. sibiricum = Ptychostomum pallens 51
B. subapiculatum = Imbribryum subapiculatum 5, 49, 50
B. tenuisetum = Imbribryum tenuisetum 5, 49, 50
B. torquescens = Ptychostomum torquescens 51
B. turbinatum = Ptychostomum turbinatum 51
B. uliginosum = Ptychostomum cernuum 50
B. valparaisense 4, 49
B. versicolor 49
B. violaceum 49
B. warneum = Ptychostomum warneum 51
B. weigelii = Ptychostomum weigelii 51
B. wrightii = Ptychostomum wrightii 51
Bucegia = Marchantia subgenus Preissia 26
B. romanica = Marchantia romanica 26
Buckia 63
B. vaucheri 63
Buckiella = Plagiothecium 55
B. undulata = Plagiothecium undulatum 56
Bucklandiella = Racomitrium subgenus Ellipticodryptodon 44
B. affinis = Racomitrium affine 44
B. elliptica = Racomitrium ellipticum 44
B. heterosticha = Racomitrium heterostichum 44
B. himalayana = Racomitrium himalayanum 44
B. lamprocarpa = Racomitrium lamprocarpum 44
87
B. lusitanica = Racomitrium lusitanicum 44
B. macounii = Racomitrium macounii 44
subsp. alpina = Racomitrium macounii subsp. alpinum 44
B. microcarpa = Racomitrium microcarpon 44
B. nivalis = Racomitrium nivale 44
B. obtusa = Racomitrium obtusum 44
B. sudetica = Racomitrium sudeticum 44
Buxbaumia 30
B. aphylla 30
B. viridis 30
BUXBAUMIACEAE 7, 30
BUXBAUMIALES 7, 30
Buxbaumiidae 7
CALLICLADIACEAE 8, 62
Callicladium 62
C. haldanianum 62
C. imponens 62
Calliergon 58
C. cordifolium 58
C. giganteum 58
C. megalophyllum 58
C. richardsonii 58
CALLIERGONACEAE 2, 8, 58
Calliergonella 63
C. cuspidata 63
C. lindbergii 63
CALOBRYALES 6, 8
[Calohypnum plumiforme] 69
Calomnion 55
C. complanatum 4, 55
Calycularia 24
C. laxa 24
Calyculariineae 7
CALYCULARIACEAE 7, 24
CALYMPERACEAE 7, 35
Calymperes 35
C. erosum 35
Calypogeia 16
C. subgenus Asperifoliae 16
C. subgenus Calypogeia 16
C. arguta 16
C. azorica 16
C. azurea 16
C. fissa 16
subsp. fissa 16
subsp. neogaea 16
var. paludosa 16
C. integristipula 16
C. muelleriana 16
[subsp. blomquistii] 16
subsp. muelleriana 16
var. erecta = Calypogeia muelleriana subsp. muelleriana 16
C. neesiana 16
subsp. neesiana 16
[subsp. subalpina] 16
var. hygrophila = Calypogeia neesiana var. neesiana 16
var. repanda = Calypogeia neesiana subsp. neesiana 16
C. sphagnicola 16
f. paludicola = Calypogeia fissa var. paludosa 16
C. suecica 16
CALYPOGEIACEAE 6, 16
Calyptrochaeta 55
C. apiculata 4, 55
Campyliadelphus 56
C. chrysophyllus 56
C. elodes 56
Campylidium = Campylophyllopsis 57
88
N. G. HODGETTS ET AL.
C. calcareum = Campylophyllopsis calcarea 57
C. sommerfeltii = Campylophyllopsis sommerfeltii 57
Campylium 56
C. bambergeri 56
C. decipiens = Drepanocladus polygamus 57
C. laxifolium 56
C. longicuspis 57
C. protensum 57
C. stellatum 57
Campylophyllopsis 57
C. calcarea 57
C. sommerfeltii 57
Campylophyllum 57
C. calcareum = Campylophyllopsis calcarea 57
C. halleri 57
C. montanum 57
C. sommerfeltii = Campylophyllopsis sommerfeltii 57
Campylopus 33
C. atrovirens 33
var. falcatus = Campylopus atrovirens 33
C. brevipilus 33
C. cygneus 33
C. flaccidus 33
C. flexuosus 33
C. fragilis 33
C. gracilis 33
C. incrassatus 33
C. introflexus 4, 33
C. oerstedianus 33
C. pilifer 33
C. pyriformis 33
var. azoricus = Campylopus pyriformis 33
C. schimperi 33
C. schwarzii = Campylopus gracilis 33
C. setifolius 33
C. shawii 33
C. subulatus 33
var. schimperi = Campylopus schimperi 33
C. subporodictyon = Atractylocarpus subporodictyon 32
Campylostelium 44
C. pitardii 44
C. saxicola 44
C. strictum 44
CATOSCOPIACEAE 7, 32
CATOSCOPIALES 7, 32
Catoscopium 32
C. nigritum 32
Cephalozia 10
C. affinis = Fuscocephaloziopsis affinis 10
C. ambigua 10
C. bicuspidata 10
subsp. bicuspidata 10
subsp. lammersiana 10
var. paludosa = Cephalozia ambigua 10
C. catenulata = Fuscocephaloziopsis catenulata 10
C. connivens = Fuscocephaloziopsis connivens 10
var. compacta = Fuscocephaloziopsis connivens subsp. connivens 10
C. crassifolia = Fuscocephaloziopsis crassifolia 11
C. crossii 10
C. lacinulata 10
C. leucantha = Fuscocephaloziopsis leucantha 11
var. robusta = Fuscocephaloziopsis leucantha 11
C. loitlesbergeri = Fuscocephaloziopsis loitlesbergeri 11
[Cephalozia lucens] 66
C. lunulifolia = Fuscocephaloziopsis lunulifolia 11
C. macounii 10
C. macrostachya = Fuscocephaloziopsis macrostachya 11
C. pleniceps = Fuscocephaloziopsis pleniceps 11
var. sphagnorum = Fuscocephaloziopsis pleniceps var. pleniceps 11
CEPHALOZIACEAE 6, 10
Cephaloziella 11
C. subgenus Cephaloziella 11
C. subgenus Dichiton 12
C. subgenus Evansia 12
C. subgenus Prionolobus 12
C. alpina = Cephaloziella varians 12
C. arctica = Cephaloziella varians 12
C. arctogena 11
C. aspericaulis 11
C. baumgartneri 11
C. calyculata 12
C. dentata 12
C. divaricata 11
var. asperifolia = Cephaloziella divaricata var. scabra 11
var. divaricata 11
var. polystratosa = Cephaloziella. polystratosa 12
var. rupestris = Cephaloziella divaricata var. divaricata 11
var. scabra 11
C. elachista 11
var. elachista 11
[var. spinophylla] 11
C. elegans 11
C. granatensis 12
C. grimsulana 11
C. hampeana 11
var. subtilis = Cephaloziella hampeana 11
C. integerrima 11
var. obtusa = Cephaloziella integerrima 12
[C. mammillifera] 66
C. massalongi 12
var. compacta = Cephaloziella massalongi 11
C. nicholsonii 11
C. phyllacantha 12
C. polystratosa 12
C. rubella 12
subsp. arctogena = Cephaloziella arctogena 11
var. bifida = Cephaloziella rubella 12
var. elegans = Cephaloziella elegans 11
var. pulchella = Cephaloziella rubella 12
var. sullivantii = Cephaloziella rubella 12
C. spinigera 12
f. striatula = Cephaloziella spinigera 12
C. stellulifera 12
var. limprichtii = Cephaloziella stellulifera 12
C. subdentata = Cephaloziella spinigera 12
C. turneri 12
C. uncinata 12
[var. brevigyna] 12
[var. mamillosa] 12
[var. sphagnicola] 12
var. uncinata 12
C. varians 12
var. arctica = Cephaloziella varians 12
var. scabra = Cephaloziella varians 12, 66
[C. verrucosa] 66
CEPHALOZIELLACEAE 6, 11
Cephaloziineae 6
Cephalozioideae 10
Ceratodon 36
C. amazonum 36
C. conicus 36
C. purpureus 36
JOURNAL OF BRYOLOGY
subsp. purpureus 36
subsp. stenocarpus 36
var. conicus = Ceratodon conicus 36
Cheilolejeunea 22
C. subgenus Euosmolejeunea 22
C. cedercreutzii 22
[C. xanthocarpa] 68
Cheilolejeuneinae (subtrib.) 22
Cheilothela 36
C. chloropus 36
Chenia 37
C. leptophylla 4, 37
C. ruigtevleia 37
Chiloscyphus 20
C. coadunatus = Lophocolea coadunata 20
C. fragilis = Chiloscyphus pallescens var. fragilis 20
C. fragrans = Lophocolea fragrans 20
C. latifolius = Lophocolea bidentata 20
C. minor = Lophocolea minor 20
C. pallescens 20
var. fragilis 20
var. pallescens 20
C. polyanthos 20
var. polyanthos 20
var. rivularis 20
C. profundus = Lophocolea heterophylla 20
C. rivularis = Chiloscyphus polyanthos var. rivularis 20
Chionoloma 40
C. bombayense 40
C. daldinianum 40
C. hibernicum 40
C. minus 41
C. recurvifolium 41
C. tenuirostre 41
var. holtii 41
var. tenuirostre 41
Cinclidium 52
C. arcticum 52
C. latifolium 52
C. minutifolium 52
C. stygium 52
C. subrotundum 52
Cinclidotus 37
[C. bistratosus] 69
C. confertus = Cinclidotus riparius 37
C. aquaticus 37
C. danubicus 37
C. fontinaloides 37
[C. pachyloma] 69
[C. pachylomoides] 68
C. riparius 37
[C. vardaranus] 69
C. vivesii 37
Cirriphyllum 60
C. crassinervium 60
C. piliferum 60
Cladopodiella = Odontoschisma 11
C. francisci = Odontoschisma francisci 11
C. fluitans = Odontoschisma fluitans 11
Claopodium 58
C. rostratum 58
C. whippleanum 58
Clasmatodon 60
C. parvulus 60
Cleistocarpidium 36
C. palustre 36
Clevea 25
C. hyalina 25
var. rufescens = Clevea hyalina 25
var. suecica = Clevea hyalina 25
C. spathysii 25
CLEVEACEAE 7, 25
CLIMACIACEAE 8, 56
Climacium 56
C. dendroides 56
Cnestrum 35
C. alpestre 35
C. glaucescens 35
C. schisti 35
Codonoblepharon 53
C. forsteri 53
Codriophorus = Racomitrium subgenus Cataracta 44
C. acicularis = Racomitrium aciculare 44
C. aquaticus = Racomitrium aquaticus 44
C. fascicularis = Racomitrium fasciculare 44
C. hespericus = Racomitrium hespericum 44
Cololejeunea 22
C. subgenus Aphanolejeunea 22
C. subgenus Cololejeunea 22
C. subgenus Diaphanae 22
C. azorica 22
C. calcarea 22
C. madeirensis 22
C. microscopica 22
[var. africana] 22
[var. exigua] 22
var. microscopica 22
C. minutissima = Myriocoleopsis minutissima 22
C. rossettiana 22
C. schaeferi 22
C. sintenisii 22
Cololejeuneinae (subtrib.) 22
Colura 22
C. subgenus Colura 22
C. section Colura 22
C. calyptrifolia 22
Conardia 57
C. compacta 57
CONOCEPHALACEAE 7, 25
Conocephalum 25
C. conicum 25
C. salebrosum 25
Conostomoideae 47
Conostomum 47
C. tetragonum 47
Corsinia 25
C. coriandrina 25
CORSINIACEAE 7, 25
Corsinioideae 25
Coscinodon 2, 44
C. cribrosus 44
C. horridus 44
C. humilis 45
C. monchiquensis 45
[C. yukonensis] 45
Cratoneuroideae 56
Cratoneuron 56
C. curvicaule 56
C. filicinum 56
Crossidium 37
C. aberrans 37
C. crassinervium 37
C. davidai 37
C. geheebii 37
89
90
N. G. HODGETTS ET AL.
C. laevipilum 37
C. laxefilamentosum 37
C. squamiferum 37
var. pottioideum 37
var. squamiferum 37
Crossocalyx 9
C. hellerianus 9
Crossogyna = Syzygiella 9
C. autumnalis = Syzygiella autumnalis 9
[Crumia latifolia] 69
Cryphaea 64
C. heteromalla 64
CRYPHAEACEAE 8, 64
Cryptoleptodon = Leptodon 64
C. longisetus = Leptodon longisetus 64
Cryptocolea 18
C. imbricata 18
Cryptothallus = Aneura 23
C. mirabilis = Aneura mirabilis 23
Ctenidium 65
C. molluscum 65
C. procerrimum = Pseudostereodon procerrimus 63
CYATHODIACEAE 7, 26
Cyathodium 26
[C. cavernarum] 66, 68
C. foetidissimum 26
[C. spurium] 66
Cyclodictyon 55
C. laetevirens 55
Cynodontium 35
C. asperifolium 35
C. bruntonii 35
C. fallax 35
C. gracilescens 35
C. jenneri 35
C. polycarpon 35
C. strumiferum 35
C. suecicum 35
C. tenellum 35
Cyrtomnium 52
C. hymenophylloides 52
C. hymenophyllum 52
Daltonia 55
D. lindigiana 55
D. splachnoides 55
D. stenophylla = Daltonia lindigiana 55
DALTONIACEAE 8, 55
Delavayelloideae 17
Dendrocryphaea 64
D. lamyana 64
Desmatodon = Tortula 39
D. bogosicus = Tortula bogosica 39
D. guepinii = Tortula guepinii 40
D. latifolius = Tortula hoppeana 40
D. laureri = Tortula laureri 40
D. leucostoma = Tortula leucostoma 40
D. meridionalis = Tortula marginata 40
D. randii = Tortula randii 40
D. systylius = Tortula systylia 40
D. ucrainicus = Tortula ucrainica 40
D. wilczekii = Hennediella heimii 38
Dialytrichia 37
D. fragilifolia = Dialytrichia saxicola 37
D. mucronata 37
D. saxicola 37
Dichelyma 55
D. capillaceum 55
D. falcatum 55
Dichiton = Cephaloziella 11
D. integerrimum = Cephaloziella integerrima 12
Dichodontium 33
D. flavescens 33
D. palustre = Diobelonella palustris 33
D. pellucidum 33
DICRANACEAE 7, 34
DICRANALES 7, 32
Dicranella 33
D. campylophylla 33
D. cerviculata 33
D. crispa 33
D. grevilleana 33
D. heteromalla 33
D. howei 33
D. humilis 33
D. palustris = Diobelonella palustris 33
D. riparia = Kiaeria riparia 35
D. rufescens 33
D. schreberiana 33
D. staphylina 33
D. subulata 33
D. varia 33
DICRANELLACEAE 7, 33
Dicranidae 7
Dicranoideae 34
Dicranodontium 33
D. asperulum 33
D. denudatum 33
var. alpinum = Dicranodontium denudatum 33
D. subporodictyon = Atractylocarpus subporodictyon 32
D. uncinatum 33
Dicranoloma 34
D. menziesii 4, 34
Dicranoweisia 35
D. cirrata 35
D. compacta = Hymenoloma compactum 32
D. crispula = Hymenoloma crispulum 32
var. compacta = Hymenoloma compactum 32
var. intermedia = Hymenoloma mulahaceni 32
D. intermedia = Hymenoloma mulahaceni 32
Dicranum 34
D. section Convolutifolia 34
D. section Crassinervia 35
D. section Dicranum 34
D. section Elongata 35
D. section Fuscescentiformia 34
D. section Montana 35
D. section Spuria 34
D. acutifolium 34
D. angustum 34
D. bardunovii 34
D. bergeri = Dicranum undulatum 34
var. acutifolium = Dicranum acutifolium 34
D. bonjeanii 34
D. brevifolium 34
D. canariense = Dicranum scottianum 35
D. congestum = Dicranum flexicaule 34
D. crassifolium 34
D. dispersum 34
D. drummondii 34
D. elongatum 35
D. flagellare 35
D. flexicaule 34
D. fragilifolium 35
D. fulvum 35
JOURNAL OF BRYOLOGY
D. fuscescens 34
var. congestum = Dicranum flexicaule 34
var. flexicaule = Dicranum flexicaule 34
D. groenlandicum 35
D. laevidens 35
D. leioneuron 34
D. majus 34
D. montanum 35
D. muehlenbeckii 34
var. acutifolium = Dicranum acutifolium 34
var. brevifolium = Dicranum brevifolium 34
var. cirrhatum = Dicranum brevifolium 34
D. polysetum 34
D. rugosum = Dicranum polysetum 34
D. schljakovii 35
D. scoparium 34
D. scottianum 35
D. septentrionale 34
D. spadiceum 35
D. spurium 34
D. strictum = Dicranum tauricum 35
D. subporodictyon = Atractylocarpus subporodictyon 32
D. tauricum 34
D. transsylvanicum 34
D. undulatum 34
D. undulatum F.Weber & D.Mohr, hom. illeg. = Dicranum polysetum 34
D. undulatum Turner, hom. illeg. = Dicranum bonjeanii 34
D. viride 35
Didymodon 5, 37
D. acutus 37
D. asperifolius 37
D. australasiae 37
D. bistratosus 37
D. brachyphyllus 37
[D. caboverdeanus] 69
D. cordatus 37
D. cylindicus var. daldianus = Chionoloma daldinianum 40
D. eckeliae 37
D. erosus = Didymodon tophaceus subsp. erosus 38
D. fallax 37
D. ferrugineus 37
D. giganteus 37
D. glaucus 5, 37
subsp. glaucus 37
subsp. verbanus 5, 37
D. icmadophilus 37
D. insulanus 37
D. johansenii 37
D. lamyanus = Didymodon brachyphyllus 37
D. luridus 37
D. maschalogenus 38
D. maximus 38
D. nicholsonii 38
D. rigidulus 38
D. sicculus = Didymodon tophaceus subsp. sicculus 38
D. sinuosus 38
D. spadiceus 38
D. subandreaeoides 38
[D. tectorum] 69
D. tomaculosus 38
D. tophaceus 38
subsp. erosus 38
subsp. sicculus 38
subsp. tophaceus 38
var. anatinus 38
var. tophaceus 38
D. umbrosus 38
D. validus 38
D. verbanus = Didymodon glaucus subsp. verbanus 37
D. vinealis 38
Dilutineuron = Racomitrium subgenus Cataracta 44
D. fasciculare = Racomitrium fasciculare 44
Diobelonella 33
D. palustris 33
DIPHYSCIACEAE 7, 30
DIPHYSCIALES 7, 30
Diphysciidae 7
Diphyscium 30
D. foliosum 30
Diplophyllum 14
D. section Diplophyllum 14
D. section Protodiplophyllum 14
D. albicans 14
D. obtusatum 14
D. obtusifolium 14
[subsp. domesticum] 14
subsp. obtusifolium 14
[D. sibiricum] 14
D. taxifolium 14
var. macrostictum = Diplophyllum taxifolium 14
[var. mucronatum] 14
var. taxifolium 14
DISCELIACEAE 7, 32
Discelium 32
D. nudum 32
DISTICHIACEAE 7, 32
Distichium 32
D. capillaceum 32
D. hagenii 32
D. inclinatum 32, 50
Distichophyllum 55
D. carinatum 55
DITRICHACEAE 7, 32, 36
Ditrichum 36
D. cornubicum 36
D. crispatissimum = Flexitrichum gracile 32
D. cylindricum = Trichodon cylindricus 36
D. flexicaule = Flexitrichum flexicaule 32
D. gracile = Flexitrichum gracile 32
D. heteromallum 36
D. lineare 36
D. pallidum 36
D. plumbicola 36
D. punctulatum 36
D. pusillum 36
D. subulatum 36
D. zonatum 36
var. scabrifolium = Ditrichum zonatum 36
Douinia 14
D. ovata 14
Drepanium 57
D. fastigiatum 57
D. recurvatum = Drepanium fastigiatum 57
Drepanocladus 57
D. aduncus 57
D. angustifolius 57
D. arcticus 57
D. brevifolius 57
D. capillifolius 57
[D. longifolius] 57, 68
D. longifolius auct. eur. = Drepanocladus capillifolius 57
D. lycopodioides 57
D. polygamus 57
91
92
N. G. HODGETTS ET AL.
D. sendtneri 57
D. sordidus 57
D. trifarius 57
D. turgescens 57
Drepanolejeunea 22
D. subgenus Drepanolejeunea 22
D. hamatifolia 22
Drepanolejeuneinae (subtrib.) 22
Dryptodon = Grimmia 45
D. orbicularis = Grimmia orbicularis 45
D. patens = Grimmia ramondii 45
Dumortiera 26
D. hirsuta 26
subsp. hirsuta 26
[subsp. nepalensis] 26
[subsp. tatunoi] 26
DUMORTIERACEAE 7, 26
ECHINODIACEAE 8, 65
Echinodium 65
E. prolixum = Isothecium prolixum 65
E. renauldii 65
E. setigerum 65
E. spinosum 65
Elodium = Helodium 59
E. blandowii = Helodium blandowii 59
Encalypta 31
E. section Encalypta 31
E. section Megasporae 31
E. section Pyromitrium 31
E. section Rhabdotheca 31
E. section Streptotheca 31
E. affinis 31
subsp. affinis 31
subsp. macounii 31
E. alpina 31
E. brevicolla 31
E. brevipes 31
E. ciliata 31
E. intermedia = Encalypta pilifera 31
E. longicolla 31
E. microstoma 31
E. mutica 31
E. obovatifolia = Encalypta pilifera 31
E. pilifera 31
E. procera 31
E. rhaptocarpa 31
var. leptodon = Encalypta trachymitra 31
var. spathulata = Encalypta spathulata 31
var. trachymitria = Encalypta trachymitria 31
E. spathulata 31
E. streptocarpa 31
E. trachymitria 31
E. vulgaris 31
ENCALYPTACEAE 7, 30
ENCALYPTALES 7, 30
Encalyptidae 7
Endogemma 16
E. caespiticia 16
ENDOGEMMATACEAE 6, 16
Entodon 64
E. challengeri 64
E. cladorrhizans 64
E. concinnus 64
[E. pseudoseductrix] 69
E. schleicheri 64
ENTODONTACEAE 8, 64
[Entodontopsis leucostega] 69
Entosthodon 31
E. subgenus Entosthodon 31
E. subgenus Murcia 31
E. subgenus Plagiodus 31
E. abramovae 31
[E. angustifolius] 69
E. attenuatus 31
E. commutatus 31
E. convexus 31
E. dagestanicus 31
E. duriaei 31
E. fascicularis 31
E. handelii 31
E. hungaricus 31
E. krausei = Entosthodon commutatus 31
E. kroonkurk 31
E. mouretii 31
E. muhlenbergii 31
[E. niloticus] 69
E. obtusus 31
E. pulchellus 31
E. schimperi 31
E. stenophyllus 31
E. templetonii = Entosthodon attenuatus 31
Ephemerella = Physcomitrium 31
E. readeri = Physcomitrium readeri 32
Ephemerum 41
E. cohaerens 41
E. crassinervium 41
subsp. rutheanum 41
subsp. sessile 41
E. hibernicum = Ephemerum crassinervium subsp.
rutheanum 41
E. minutissimum = Ephemerum serratum 41
E. recurvifolium 41
E. rutheanum = Ephemerum crassinervium subsp.
rutheanum 41
E. serratum 41
var. minutissimum = Ephemerum serratum 41
var. rutheanum = Ephemerum crassinervium subsp. rutheanum 41
E. serratum auct. = Ephemerum stoloniferum 41
E. sessile = Ephemerum crassinervium subsp. sessile 41
E. spinulosum 41
E. stellatum = Ephemerum stoloniferum 41
E. stoloniferum 41
Epipterygium 52
[E. rigidum] 69
E. tozeri 52
Eremonotus 17
E. myriocarpus 17
[Erpodium grossirete] 69
[Erpodium perrottetii] 69
Eucladium 41
E. verbanum = Didymodon glaucus subsp. verbanum 5
E. verticillatum 41
var. angustifolium = Eucladium verticillatum 41
Eurhynchiastrum 61
E. diversifolium 61
E. pulchellum 61
var. praecox = Eurhynchiastrum pulchellum 61
var. diversifolium = Eurhynchiastrum diversifolium 61
Eurhynchioideae 59
Eurhynchium 59
E. angustirete 59
E. striatum 59
Exobryum = Didymodon 37
JOURNAL OF BRYOLOGY
E. asperifolium = Didymodon asperifolius 37
E. johansenii = Didymodon johansenii 37
Exormotheca 26
E. subgenus Corbierella 26
E. subgenus Exormotheca 26
E. pustulosa 26
E. welwitschii 26, 67
EXORMOTHECACEAE 7, 26
Exsertotheca 64
E. baetica 64
E. crispa 64
E. intermedia 64
Fabronia 56
F. altaica 56
F. ciliaris 56
[F. gueinzii] 69
[F. leikipiae] 69
F. major 56
F. pusilla 56
FABRONIACEAE 8, 56
[Fimbraria = Asterella] 66
[F. blumeana = Asterella blumeana] 66
[F. elegans = Asterella elegans] 66
[F. marginata = Asterella marginata] 66
[F. raddii = Asterella wallichiana] 66
[F. stahlii = Asterella pringlei] 66
[F. tenella = Asterella tenella] 66
Fissidens 5, 33
F. subgenus Aloma 34
F. subgenus Fissidens 34
F. subgenus Octodiceras 34
F. subgenus Pachyfissidens 5, 33
F. section Amblyothallia 34
F. section Pachyfissidens 5, 33
F. adianthoides 33
F. algarvicus = Fissidens curvatus 34
[F. allorgei] 69
[F. androgynus] 69
F. arcticus 34
F. arnoldii 34
F. asplenioides 34
F. azoricus 33
F. bambergeri = Fissidens viridulus 34
F. bryoides 34
var. bryoides 34
var. caespitans 34
var. curnovii = Fissidens bryoides var. caespitans 34
F. celticus 34
F. coacervatus 34
F. crassipes 34
subsp. crassipes 34
subsp. warnstorfii 34
F. crispus 34
F. cristatus = Fissidens dubius 33
F. curnovii = Fissidens bryoides var. caespitans 34
F. curvatus 34
[F. dankelmannii] 69
F. dubius 33
var. dubius 33
var. mucronatus 33
F. exiguus auct. eur. = Fissidens viridulus 34
F. exilis 34
[F. flaccidus] 69
F. fontanus 34
F. gracilifolius 34
F. grandifrons 33
F. gymnandrus 34
F. herzogii = Fissidens crispus 34
F. incurvus 34
F. jansenii 34
F. limbatus = Fissidens crispus 34
F. luisieri = Fissidens serrulatus 34
[F. megalotis subsp. helictocaulos] 69
F. microstictus 34
F. minutulus = Fissidens crispus 34
F. monguillonii 34
F. nobreganus 34
F. osmundoides 33
F. ovatifolius 34
F. polyphyllus 33
F. pusillus 34
F. rivularis 34
F. rufulus 34
[F. sciophyllus] 69
F. serratus 34
F. serrulatus 33
F. sublimbatus 34
F. sublineaefolius 34
F. taxifolius 33
subsp. pallidicaulis = Fissidens taxifolius 33
[F. usambaricus] 69
F. viridulus 34
var. bambergeri = Fissidens viridulus 34
var. incurvus = Fissidens incurvus 34
var. pusillus = Fissidens pusillus 34
var. tenuifolius = Fissidens gracilifolius 34
FISSIDENTACEAE 2, 7, 33
FLEXITRICHACEAE 7, 32
Flexitrichum 32
F. flexicaule 32
F. gracile 32
FONTINALACEAE 8, 55
Fontinalis 55
F. antipyretica 55
subsp. antipyretica 55
subsp. bryhnii 55
subsp. gracilis 55
subsp. kindbergii 55
[var. heldreichii] 69
F. dalecarlica 55
F. dichelymoides 55
F. hypnoides 55
var. duriaei 55
var. hypnoides 55
F. squamosa 55
var. curnowii = Fontinalis squamosa 55
var. dixonii = Fontinalis squamosa 55
Fossombronia 24
F. angulosa 24
F. caespitiformis 24
subsp. caespitiformis 24
subsp. multispira 24
F. crispa 24
F. crozalsii = Fossombronia mittenii 24
F. echinata 24
F. fimbriata 24
F. fleischeri 24
F. foveolata 24
F. husnotii = Fossombronia caespitiformis subsp.
multispira 24
F. incurva 24
F. leucoxantha 24
F. maritima 24
F. mittenii 24
93
94
N. G. HODGETTS ET AL.
F. pusilla 24
F. wondraczekii 24
F. zeyheri 24
FOSSOMBRONIACEAE 7, 24
FOSSOMBRONIALES 6, 24
Fossombroniineae 7
Frullania 21
F. subgenus Frullania 21
F. subgenus Thyopsiella 21
F. acicularis 21
F. azorica 21, 67
F. bolanderi 22
F. calcarifera 22
F. cesatiana = Frullania riparia 21, 67
[var. muscicola = Frullania muscicola] 21
F. cleistostoma 22
[F. davurica] 66
subsp. jackii = Frullania jackii 21, 66
F. dilatata 21
[subsp. asiatica] 21
subsp. dilatata 21
F. ericoides 21
var. ericoides 21
[var. laxa] 21
[var. minor] 21
[var. verrucosa] 21
F. fragilifolia 21
F. inflata auct. = F. cleistostoma 22
[F. inflata] 22, 66, 67
F. jackii 21, 66
F. microphylla 21
var. deciduifolia = Frullania microphylla 21
[Frullania muscicola] 21, 67
F. oakesiana 21
subsp. oakesiana 21
[subsp. takayuensis] 21
[F. obscurifolia] 67
F. parvistipula 21
F. polysticta 21
F. riparia 21
F. sergiae 21
[F. socotrana] 68
[F. spongiosa] 68
F. stylifera 21
F. subarctica 21
F. tamarisci 21
var. atrovirens = Frullania tamarisci 21
var. azorica = Frullania acicularis 21
var. cornubica = Frullania tamarisci 21
var. ericetorum = Frullania tamarisci 21
var. robusta = Frullania tamarisci 21
var. sardoa = Frullania tamarisci 21
F. teneriffae 21
FRULLANIACEAE 6, 21
Funaria 31
[F. anomala] 4
[F. altissima] 69
[F. chevalieri] 69
F. dentata = Entosthodon muhlenbergii 31
F. handelii = Entosthodon handelii 31
F. aequidens 31
F. arctica 31
F. hygrometrica 31
F. microstoma 31
FUNARIACEAE 2, 7, 31
FUNARIALES 7, 31
Funariidae 7
Funariella 31
F. curviseta 31
Funarioideae 31
Fuscobryum = Didymodon 37
F. subandreaeoides = Didymodon subandreaeoides 38
Fuscocephaloziopsis 10
F. affinis 10
F. albescens 10
var. albescens 10
var. islandica 10
F. catenulata 10
subsp. catenulata 10
[subsp. nipponica] 10
F. connivens 10
subsp. connivens 10
[subsp. fissa] 10
[subsp. sandwicensis] 10
F. crassifolia 11
F. leucantha 11
F. loitlesbergeri 11
F. lunulifolia 11
F. macrostachya 11
[subsp. australis] 11
subsp. macrostachya 11
var. macrostachya 11
var. spiniflora 11
F. pleniceps 11
[var. caroliniana] 11
var. pleniceps 11
Geheebia = Didymodon 37
G. fallax = Didymodon fallax 37
G. ferruginea = Didymodon ferrugineus 37
G. gigantea = Didymodon giganteus 37
G. maschalogena = Didymodon maschalogenus 38
G. maxima = Didymodon maximus 38
G. spadicea = Didymodon spadiceus 38
G. tophacea = Didymodon tophaceus 38
Gemmabryum = Bryum 3, 48
G. apiculatum = Anomobryum apiculatum 48
G. barnesii = Bryum dichotomum 48
G. caespiticium = Ptychostomum imbricatulum 50
G. demaretianum = Bryum demaretianum 48
G. dichotomum = Bryum dichotomum 49
G. gemmiferum = Bryum gemmiferum 49
G. gemmilucens = Bryum gemmilucens 49
G. klinggraeffii = Bryum klinggraeffii 49
G. kunzei = Ptychostomum kunzei 50
G. radiculosum = Bryum radiculosum 49
G. ruderale = Bryum ruderale 49
G. subapiculatum = Imbribryum subapiculatum 49
G. tenuisetum = Imbribryum tenuisetum 49
G. valparaisense = Bryum valparaisense 49
G. violaceum = Bryum violaceum 49
GEOCALYCACEAE 6, 16
Geocalyx 16
G. graveolens 16
GIGASPERMACEAE 7, 32
GIGASPERMALES 7, 32
Gigaspermidae 7
Gigaspermum 32
G. mouretii 32
Glyphomitrium 35
G. daviesii 35
Gongylanthus 19
G. ericetorum 19
JOURNAL OF BRYOLOGY
Goniomitrium 31
G. seroi 31
Grimmia 2, 45
G. alpestris 45
G. anodon 45
G. anomala 45
G. arenaria 45
G. atrata 45
G. britannica = Grimmia trichophylla 45
G. caespiticia 45
G. capillata 45
G. cribrosa = Coscinodon cribrosus 44
G. crinita 45
G. crinitoleucophaea 45
G. curvata = Grimmia ramondii 45
G. curviseta 45
G. decipiens 45
G. dissimulata 45
G. donniana 45
G. elatior 45
G. elongata 45
G. funalis 45
G. fuscolutea 45
G. hartmanii 45
G. horrida = Coscinodon horridus 44
G. incurva 45
G. laevigata 45
G. lisae 45
G. longirostris 45
G. meridionalis 45
G. meteorae = Grimmia nutans 45
G. mollis 45
G. montana 45
G. muehlenbeckii 45
G. nutans 45
G. orbicularis 45
G. ovalis 45
G. pitardii = Campylostelium pitardii 44
G. plagiopodia 45
G. poecilostoma = Grimmia crinitoleucophaea 45
G. pulvinata 45
G. ramondii 45
G. reflexidens 45
G. sessitana = Grimmia reflexidens 45
G. teretinervis 45
G. tergestina 45
var. tergestinoides = Grimmia tergestina 45
G. torquata 45
G. trichophylla 45
var. meridionalis = Grimmia meridionalis 45
G. triformis 45
G. ungeri 45
G. unicolor 45
GRIMMIACEAE 7, 44
GRIMMIALES 7, 32, 43
Grimmioideae 44
[Groutiella tomentosa] 69
Guembelia = Grimmia 45
G. laevigata = Grimmia laevigata 45
G. longirostris = Grimmia longirostris 45
G. ovalis = Grimmia ovalis 45
G. tergestina = Grimmia tergestina 45
Gymnobarbula 36, 41
G. bicolor 41
Gymnocolea 9
G. acutiloba = Gymnocolea inflata subsp. acutiloba 9
G. borealis 9
G. fascinifera 9
G. inflata 9
subsp. acutiloba 9
subsp. inflata 9
var. heterostipa = Gymnocolea inflata subsp. inflata 9
GYMNOMITRIACEAE 6, 16
Gymnomitrioideae 16
Gymnomitrion 16
G. adustum 16
G. alpinum 16
G. apiculatum = Marsupella apiculata 16
G. brevissimum 16
G. commutatum 16
G. concinnatum 16
G. corallioides 16
G. crenulatum 16
G. obtusum 16
G. revolutum 16
[subsp. novoguineanense] 16
subsp. revolutum 16
[Gymnostomiella erosula] 69
[Gymnostomiella vernicosa] 69
Gymnostomum 41
G. aeruginosum 41
var. aeruginosum 41
var. obscurum 41
G. boreale = Hymenostylium gracillimum 42
G. calcareum 41
var. lanceolatum = Gymnostomum calcareum 41
var. atlanticum 41
var. calcareum 41
G. lanceolatum = Gymnostomum calcareum 41
G. recurvirostrum = Hymenostylium recurvirostrum 42
G. viridulum 41
Gyroweisia 41
G. reflexa 41
G. tenuis 41
Habrodon 56
H. perpusillus 56
HABRODONTACEAE 8, 56
Hageniella 63
H. micans 63
Hamatocaulis 58
H. lapponicus 58
H. vernicosus 58
Haplocladium 59
H. angustifolium 59
H. microphyllum 59
H. virginianum 59
Haplohymenium = Anomodon 66
H. triste = Anomodon tristis 66
HAPLOMITRIACEAE 6, 8
Haplomitriidae 6
Haplomitriopsida 6, 8
Haplomitrium 8
H. subgenus Haplomitrium 8
H. section Haplomitrium 8
H. hookeri 8
var. hookeri 9
[var. minutum] 9
Harpalejeunea 22
H. subgenus Harpalejeunea 22
H. molleri 22
[subsp. integra] 22
subsp. molleri 22
HARPANTHACEAE 6, 17
Harpanthus 17
95
96
N. G. HODGETTS ET AL.
H. flotovianus 17
var. chiloscyphoides = Harpanthus flotovianus 17
var. latifolia = Harpanthus flotovianus 17
var. retusa = Harpanthus flotovianus 17
H. scutatus 17
Hedenasiastrum 60
H. percurrens 60
Hedwigia 47
H. ciliata 47
var. leucophaea = Hedwigia emodica 47
H. emodica 47
[H. integrifolia] 47
H. integrifolia = Braunia imberbis 47
H. mollis 47
H. nemoralis 47
H. stellata 47
H. striata 47
HEDWIGIACEAE 7, 46
HEDWIGIALES 7, 46
Hedwigidium = Braunia 46
H. integrifolium = Braunia imberbis 47
Helicodontium 60
H. capillare 60
Helicodontoideae 60
Helodium 59
H. blandowii 59
Hennediella 38
H. heimii 38
var. arctica 38
var. heimii 38
H. macrophylla 4, 38
H. stanfordensis 4, 38
HERBERTACEAE 6, 19
Herbertus 19
[Herbertus aduncus] 19, 116
subsp. hutchinsiae = H. hutchinsiae 19
H. aduncus auct eur = Herbertus hutchinsiae 19
H. azoricus 19
H. borealis 19, 67, 116
H. delavayi auct. = Herbertus borealis 19
[H. delavayi] 19
[H. dicranus] 67
H. hutchinsiae 19, 116
[H. juniperoideus] 19
H. norenus 19, 116
[H. sakuraii] 67
H. sendtneri 19
H. stramineus 19, 116
[H. tenuis] 19
[Herpetineuron toccoae] 69
Herzogiella 55
H. seligeri 55
H. striatella 55
H. turfacea 55
Heterocladiella 65
H. dimorpha 65
HETEROCLADIELLACEAE 8, 65
Heterocladium 65
H. dimorphum = Heterocladiella dimorpha 65
H. flaccidum 65
H. heteropterum 65
H. wulfsbergii 65
Heterogemma 12
H. capitata 12
H. laxa 12
Heterophyllium 62
H. affine = Heterophyllium nemorosum 62
H. nemorosum 62
Heteroscyphus 20
H. denticulatus 20
H. fissistipus 4, 20
[var. fissistipus] 20
[var. multispinus] 20
[var. repandus] 20
[H. integrifolius] 67
Hilpertia 38
H. velenovskyi 38
Homalia 64
H. lusitanica 64
H. trichomanoides 64
H. webbiana = Pseudomalia webbiana 65
Homalothecium 61
H. aureum 61
H. lutescens 61
var. fallax 61
var. lutescens 61
H. mandonii 61
H. meridionale 61
H. philippeanum 61
H. sericeum 61
Homomallium 63
H. incurvatum 63
Hookeria 55
[H. acutifolia] 69
H. lucens 55
HOOKERIACEAE 8, 55
HOOKERIALES 2, 8, 55
Hydrogonium 36, 41
H. amplexifolium 41
[H. arcuatum] 69
H. bolleanum 41
H. consanguineum 41
[var. cancellatum] 41
[var. consanguineum] 41
var. kurilense 41, 68
H. croceum 41
[H. orientale] 68, 69
Hydrogrimmia = Grimmia 45
H. mollis = Grimmia mollis 45
Hygroamblystegium 57
H. fluviatile 57
H. humile 57
H. tenax 57
H. varium 57
var. humile = Hygroamblystegium humile 57
Hygrobiella 17
H. laxifolia 17
HYGROBIELLACEAE 6, 17
Hygrohypnella 58
H. ochracea 58
H. polaris 58
Hygrohypnum 57
H. alpestre = Platyhypnum alpestre 57
H. alpinum = Platyhypnum alpinum 57
H. cochlearifolium = Platyhypnum cochlearifolium 58
H. duriusculum = Platyhypnum duriusculum 58
H. eugyrium = Pseudohygrohypnum eugyrium 63
H. luridum 57
H. molle = Platyhypnum molle 58
H. montanum = Campylophyllum montanum 57
H. norvegicum = Platyhypnum norvegicum 58
H. ochraceum = Hygrohypnella ochracea 58
H. polare = Hygrohypnella polaris 58
H. smithii = Platyhypnum smithii 58
JOURNAL OF BRYOLOGY
H. styriacum 57
H. subeugyrium = Pseudohygrohypnum subeugyrium 63
HYLOCOMIACEAE 8, 63
Hylocomiadelphus 63
H. triquetrus 63
Hylocomiastrum 63
H. pyrenaicum 63
H. umbratum 63
Hylocomium 63
H. splendens 63
Hymenoloma 32
H. compactum 32
H. crispulum 32
H. intermedium = H. mulahaceni 32
H. mulahaceni 32
HYMENOLOMATACEAE 7, 32
Hymenostylium 42
[H. congoanum] 69
H. gracillimum 42
[H. hildebrandtii] 69
H. insigne = Hymenostylium recurvirostrum var. insigne 42
H. recurvirostrum 42
var. insigne 42
var. recurvirostrum 42
H. xerophilum 42
Hyocomium 65
H. armoricum 65
Hyophila 42
H. involuta 42
HYPNACEAE 8, 62
HYPNALES 8, 55
Hypnelloideae 55
Hypnum 62
H. aemulans = Stereodon aemulans 63
H. adfine = Heterophyllium nemorosum 62
H. affine = Heterophyllium nemorosum 62
H. andoi 62
H. bambergeri = Campylium bambergeri 56
H. boreale = Ptychostomum pallescens 51
H. callichroum = Stereodon callichrous 63
H. cupressiforme 62
var. cupressiforme 62
var. filiforme 62
var. heseleri 62
var. lacunosum 62
var. resupinatum = Hypnum resupinatum 62
var. subjulaceum 62
H. fertile = Pseudohygrohypnum fertile 63
H. hamulosum = Stereodon hamulosus 63
H. holmenii = Stereodon holmenii 63
H. imponens = Callicladium imponens 62
H. jutlandicum 62
H. nemorosum = Heterophyllium nemorosum 62
H. pallescens = Jochenia pallescens 62
var. protuberans = Jochenia protuberans 62
var. reptile = Jochenia pallescens 62
H. plicatulum = Aquilonium plicatulum 63
H. procerrimum = Pseudostereodon procerrimus 63
H. protuberans = Jochenia protuberans 62
H. recurvatum = Drepanium fastigiatum 57
H. reptile = Jochenia pallescens 62
H. resupinatum 62
H. revolutum = Roaldia revoluta 63
var. dolomiticum = Roaldia dolomitica 63
H. sauteri = Microhypnum sauteri 57
H. subcomplanatum = Hypnum cupressiforme var.
subjulaceum 62
97
H. subimponens = Stereodon subimponens 63
H. subjulaceum = Hypnum cupressiforme var. subjulaceum 62
H. uncinulatum 62
H. vaucheri = Buckia vaucheri 63
HYPOPTERYGIACEAE 2, 8, 55
HYPOPTERYGIALES = HOOKERIALES 2, 55
Hypopterygium 2, 55
H. tamarisci 4, 55
Imbribryum 3, 49
I. alpinum 49
I. gemmiparum = Bryum gemmiparum 49
I. mildeanum 49
I. miniatum 49
I. muehlenbeckii 49
I. subapiculatum 5, 49
I. tenuisetum 5, 49
Indusiella 44
I. thianschanica 44
Isopaches 9
I. alboviridis 9
I. bicrenatus 9
I. decolorans 9
Isopterygiopsis 55
I. alpicola 55
I. muelleriana 55
I. pulchella 55
Isopterygium 62
I. tenerum 62
Isothecium 65
I. algarvicum 65
I. alopecuroides 65
I. holtii 65
I. interludens 65
I. montanum 65
I. myosuroides 65
subsp. brevinerve = Isothecium myosuroides 65
var. brachythecioides = Isothecium interludens 65
I. prolixum 65
Iwatsukiella 64
I. leucotricha 64
Jaffueliobryum 44
J. latifolium 44
Jamesoniella = Syzygiella 9
J. autumnalis = Syzygiella autumnalis 9
J. rubricaulis = Syzygiella rubricaulis 9
J. undulifolia = Biantheridion undulifolium 9
Jamesonielloideae 9
Jochenia 62
J. pallescens 62
J. protuberans 62
JOCHENIACEAE 8, 62
Jubula 22
J. hutchinsiae 22
[subsp. australiae] 22
[subsp. bogotensis] 22
subsp. caucasica 22, 67
subsp. hutchinsiae 22
[subsp. japonica] 22
[subsp. javanica] 22, 67
[subsp. pennsylvanica] 22, 67
JUBULACEAE 6, 22
Jubulineae 6
Jungermannia 17, 66
J. atrovirens 17
J. borealis 17
J. caespiticia = Endogemma caespiticia 16
J. calcicola 17
98
N. G. HODGETTS ET AL.
J. callithrix = Solenostoma callithrix 18
J. confertissima = Solenostoma confertissimum 18
J. eucordifolia 17, 67
[J. exsertifolia] 67
subsp. cordifolia = Jungermannia eucordifolia 17, 67
J. gracillima = Solenostoma gracillimum 18
var. crenulata = Solenostoma gracillimum 18
J. handelii = Solenostoma handelii 18
J. hyalina = Solenostoma hyalinum 18
J. jenseniana = Solenostoma sphaerocarpum 18
J. lanceolata auct. = Liochlaena lanceolata 17
J. lanceolata = Jungermannia atrovirens 17
var. atrovirens = Jungermannia atrovirens 17
J. leiantha = Liochlaena lanceolata 17
J. obovata = Solenostoma obovatum 18
subsp. minor = Solenostoma subellipticum 18
J. paroica = Solenostoma paroicum 18
J. polaris 17
J. pumila 17
subsp. polaris = Jungermannia polaris 17
var. alpestris = Jungermannia pumila 17
J. sphaerocarpa = Solenostoma sphaerocarpum 18
var. nana = Solenostoma sphaerocarpum 18
J. subelliptica = Solenostoma subellipticum 18
J. subulata = Liochlaena subulata 17
var. leiantha = Liochlaena lanceolata 17
JUNGERMANNIACEAE 6, 17
Jungermanniales 6, 9
Jungermanniidae 6
Jungermanniineae 6
Jungermannioideae 17
Jungermanniopsida 6, 9
Kiaeria 35
K. blyttii 35
K. falcata 35
K. glacialis 35
K. riparia 35
K. starkei 35
Kindbergia 61
K. praelonga 61
Kurzia 19
[K. makinoana] 67
K. pauciflora 19
K. sylvatica 19, 67
K. trichoclados 19
Leiocolea = Mesoptychia 17
L. alpestris = Mesoptychia collaris 18
L. badensis = Mesoptychia badensis 17
L. bantriensis = Mesoptychia bantriensis 18
subsp. wallfischii = Mesoptychia bantriensis subsp.
wallfischii 18
L. collaris = Mesoptychia collaris 18
L. fitzgeraldiae = Mesoptychia fitzgeraldiae 18
L. gillmanii = Mesoptychia gillmanii 18
L. heterocolpos = Mesoptychia heterocolpos 18
L. rutheana = Mesoptychia rutheana 18
var. laxa = Mesoptychia gillmanii var. laxa 18
L. turbinata = Mesoptychia turbinata 18
Lejeunea 22
L. subgenus Lejeunea 22
L. subgenus Nanolejeunea 22
L. canariensis 22, 67
[L. capensis] 68
L. cavifolia 22
[L. diversifolia = Microlejeunea diversifolia] 67
[L. diversiloba = Microlejeunea diversifolia] 67
L. eckloniana 22
L. flava 22
[subsp. flava] 22
subsp. moorei 22
[subsp. orientalis] 22
[subsp. tabularis] 22
[var. pellucida] 22
L. hibernica 22
L. holtii = Lejeunea eckloniana 22
[L. laetevirens] 22, 67
L. laetevirens auct. = Lejeunea canariensis 22
L. lamacerina 22
[subsp. gemminata] 22
subsp. lamacerina 22
L. mandonii 22
L. patens 22
L. ulicina = Microlejeunea ulicina 23
LEJEUNEACEAE 6, 8, 22
Lejeuneeae (trib.) 22
Lejeuneinae (subtrib.) 22
Lejeuneoideae 22
Lembidioideae 19
LEMBOPHYLLACEAE 8, 65
Lepidozia 19
L. azorica = Telaranea azorica 19
L. cupressina 19
[subsp. africana] 19
subsp. cupressina 19
[subsp. natalensis] 19
subsp. pinnata = Lepidozia cupressina subsp. cupressina 19
var. dissitifolia = Lepidozia cupressina subsp. cupressina 19
L. pearsonii 19
L. reptans 19
var. julacea = Lepidozia reptans 19
var. tenera = Lepidozia reptans 19
L. stuhlmannii 19
[subsp. pulvinata] 19
[var. carnosa] 19
LEPIDOZIACEAE 6, 19
Lepidozioideae 19
Leptobarbula 42
L. berica 42
Leptobryum 48
L. pyriforme 48
Leptodictyum 57
[L. kurdicum] 69
L. riparium 57
Leptodon 64
L. corsicus 64
L. longisetus 64
L. smithii 64
Leptodontium 38
L. flexifolium 38
L. gemmascens 38
L. proliferum 4, 38
L. styriacum 38
Leptophascum = Chenia 37
L. leptophyllum = Chenia leptophylla 37
[Leptostomopsis] 48
Leptotheca 55
L. gaudichaudii 4, 55
Leptoscyphus 20
L. subgenus Anomylia 20
L. subgenus Leptoscyphus 20
L. section Leptoscyphus 20
L. azoricus = L. porphyrius subsp. azoricus 20
JOURNAL OF BRYOLOGY
L. cuneifolius 20
L. porphyrius 20
subsp. azoricus 20
[subsp. porphyrius] 20
Lescuraea 59
L. incurvata 59
L. mutabilis 59
L. patens 59
L. plicata 59
L. radicosa 59
L. saviana 59
L. saxicola 59
L. secunda 59
Leskea 59
L. polycarpa 59
LESKEACEAE 8, 58
LEUCOBRYACEAE 7, 32
Leucobryum 33
L. albidum 33
L. glaucum 33
L. juniperoideum 33
Leucodon 64
[L. bowringii] 69
L. canariensis 64
[L. coreensis] 69
L. flagellaris 64
L. immersus 64
L. pendulus 64
L. sciuroides 64
var. morensis = Leucodon sciuroides 64
L. treleasei 64
LEUCODONTACEAE 8, 64
LEUCOMIACEAE 8, 55
Lewinskya 53
L. acuminata 53
L. affinis 53
L. breviseta 53
L. elegans 53
L. fastigiata 53
L. iberica 53
L. iwatsukii 53
L. laevigata 53
L. lamyana 53
L. pylaisii 53
L. rupestris 53
L. shawii 53
L. sordida 53
L. speciosa 53
L. striata 53
L. tortidontia 53
L. transcaucasica 53
L. vladikavkana 53
Lindbergia 59
[L. brachyptera] 68
L. brachyptera auct. eur. = Lindbergia grandiretis 59
L. dagestanica 59
L. grandiretis 59
Liochlaena 17
L. lanceolata 17
L. subulata 17
Loeskeobryum 64
L. brevirostre 64
Loeskypnum 58
L. badium 58
Lophocolea 20
L. bidentata 20
var. rivularis = Lophocolea coadunata 20
99
L. bispinosa 4, 20
L. brookwoodiana 4, 20
L. coadunata 20
L. cuspidata = L. bidentata 20
L. cuspidata auct. = L. coadunata 20
L. fragrans 20
[subsp. cocosana] 20
subsp. fragrans 20
L. heterophylla 20
[subsp. cladogyna] 20
subsp. heterophylla 20
[L. humistrata] 67
L. minor 20
L. semiteres 4, 20
[var. retusa] 20
var. semiteres 20
LOPHOCOLEACEAE 6, 20
Lophocoleineae 6
Lophozia 12
L. alpestris auct. = Barbilophozia sudetica 9
L. alpestris = Mesoptychia collaris 18
var. libertae = Mesoptychia collaris 18
L. ascendens 12
L. atlantica = Orthocaulis atlanticus 10
L. attenuata = Neoorthocaulis attenuatus 9
L. badensis = Mesoptychia badensis 17
var. obtusiloba = Mesoptychia badensis 17
L. bantriensis = Mesoptychia bantriensis 18
var. subcompressa = Mesoptychia bantriensis subsp. bantriensis 18
L. barbata = Barbilophozia barbata 9
L. bicrenata = Isopaches bicrenatus 9
L. binsteadii = Neoorthocaulis binsteadii 10
L. capitata = Heterogemma capitata 12
L. cavifolia = Orthocaulis cavifolius 10
L. ciliata 12
L. confertifolia = Lophozia wenzelii 13
L. confertifolia [sensu Schljakov 1975, 1998] = Lophozia ventricosa 13
L. confertifolia (sensu Damsholt 2002; Köckinger 2017) =
Lophozia wenzelii 13
L. confertifolia [sensu Konstantinova et al. 1992] = Lophozia
murmanica 12
L. debiliformis = Barbilophozia sudetica 9
var. concolor = Barbilophozia sudetica 9
L. decolorans = Isopaches decolorans 9
L. elongata = Protolophozia elongata 12
L. excisa = Lophoziopsis excisa 13
var. cylindracea = Lophoziopsis excisa var. excisa 13
L. floerkei = Neoorthocaulis floerkei 10
L. fuscovirens 12
L. gillmanii = Mesoptychia gillmanii 18
var. acutifolia = Mesoptychia gillmanii 18
L. gracillima = Lophozia ascendens 12
L. grandiretis = Schistochilopsis grandiretis 15
var. parviretis = Schistochilopsis grandiretis 15
var. proteidea = Schistochilopsis grandiretis 15
L. groenlandica [sensu Grolle and Long 2000; Söderström et al.
2002] = Lophozia murmanica 12
L. groenlandica [sensu Schuster 1969] = Lophozia
schusteriana 13
L. groenlandica [sensu Schljakov 1980, 1998] = Lophozia
ventricosa 13
L. groenlandica [sensu Damsholt 1994] = Lophozia wenzelii 13
L. guttulata 12
L. hatcheri = Barbilophozia hatcheri 9
L. herzogiana = Protolophozia herzogiana 12
100
N. G. HODGETTS ET AL.
L. heterocolpos = Mesoptychia heterocolpos 18
var. arctica = Mesoptychia heterocolpos var. arctica 18
var. harpanthoides = Mesoptychia heterocolpos var. harpanthoides 18
L. heteromorpha = Lophozia murmanica 13
L. hyperarctica = Schistochilopsis hyperarctica 15
L. incisa = Schistochilopsis incisa 15
subsp. opacifolia = Schistochilopsis opacifolia 15
L. jurensis = Lophoziopsis jurensis 13
L. iremelensis = Lophozia wenzelii 13
L. kunzeana = Schljakovia kunzeana 10
[L. lacerata] 69
L. lantratoviae 12
L. latifolia = Lophoziopsis jurensis 13
L. laxa = Heterogemma laxa 12
L. longidens = Lophoziopsis longidens 13
L. longiflora auct. = Lophozia guttulata 12
L. longiflora 12
var. guttulata = Lophozia guttulata 12
L. lycopodioides = Barbilophozia lycopodioides 9
L. murmanica 12
L. murmanica [sensu Schljakov 1970] = Lophozia
ventricosa 13
L. murmanica [sensu Schljakov 1969] = Lophozia savicziae 13
L. obtusa = Obtusifolium obtusum 12
L. opacifolia = Schistochilopsis opacifolia 15
L. perssonii = Oleolophozia perssonii 12
L. pellucida = Lophoziopsis pellucida 13
L. polaris = Lophoziopsis polaris 14
L. porphyroleuca = Lophozia guttulata 12
L. propagulifera auct. eur. = Lophoziopsis jurensis 13
L. quadriloba = Schljakovianthus quadrilobus 10
var. glareosa = Schljakovianthus quadrilobus 10
L. rubescens = Barbilophozia rubescens 9
L. rubrigemma = Lophoziopsis rubrigemma 14
L. rutheana = Mesoptychia rutheana 18
var. laxa = Mesoptychia gillmannii var. laxa 18
L. savicziae 13
L. schultzii = Mesoptychia rutheana 5
var. laxa = Mesoptychia gillmannii var. laxa 5
L. schusteriana 13
L. silvicola 13
L. silvicola var. grandiretis = Lophozia savicziae 13
L. silvicoloides 13
L. sudetica = Barbilophozia sudetica 9
var. anomala = Barbilophozia sudetica 9
L. subapiculata 13
L. ventricosa auct. = Lophozia silvicola 13
L. ventricosa 13
var. confusa = Lophozia ventricosa 13
var. grandiretis = Lophozia savicziae 13
var. longiflora = Lophozia longiflora 12
var. silvicola = Lophozia silvicola 13
var. uliginosa = Lophozia wenzelii 13
var. uliginosa auct. [sensu Söderström et al. 2002] =
Lophozia longiflora 12
L. wenzelii 13
var. groenlandica [sensu Konstantinova et al. 2009] =
Lophozia murmanica 13
var. lapponica 13
var. litoralis 13
var. massularioides 13
var. wenzelii 13
LOPHOZIACEAE 6, 12
Lophoziopsis 6, 13
L. excisa 13
var. elegans 13
var. excisa 13
[var. infuscata] 13, 69
[var. succulenta] 13, 69
L. jurensis 13
L. latifolia = Lophoziopsis jurensis 13
L. longidens 13
subsp. arctica 13
subsp. longidens 13
L. pellucida 13
var. minor 14
var. pellucida 14
L. perssonii = Oleolophozia perssonii 12
L. polaris 14
var. polaris 14
var. sphagnorum 14
[L. propagulifera] 13, 67
L. propagulifera auct. eur. = Lophoziopsis jurensis 13
L. rubrigemma 14
f. arctica = Lophoziopsis longidens var. arctica 13
[Luisierella] 32
Lunularia 25
L. cruciata 25
subsp. cruciata 25
[subsp. thaxteri] 25
LUNULARIACEAE 7, 25
LUNULARIALES 7, 25
Mannia 25
M. subgenus Mannia 25
M. subgenus Neesiella 25
M. androgyna 25
M. californica 25
M. controversa 25
[subsp. asiatica] 25
subsp. controversa 25
M. fragrans 25
subsp. fragrans 25
[subsp. orientalis] 25
var. brevipes = Mannia fragrans subsp. fragrans 25
M. gracilis 25
M. pilosa 25
M. sibirica 25
M. triandra 25
Marchantia 26
M. subgenus Chlamidium 26
M. subgenus Marchantia 26
M. subgenus Preissia 26
M. section Paleacea 26
M. alpestris = Marchantia polymorpha subsp.
montivagans 26
M. aquatica = Marchantia polymorpha subsp.
polymorpha 26
[M. debilis] 69
M. latifolia = Marchantia polymorpha subsp. ruderalis 26
M. paleacea 26
[subsp. diptera] 26
subsp. paleacea 26
[M. pappeana] 69
M. polymorpha 26
subsp. montivagans 26
subsp. polymorpha 26
subsp. ruderalis 26
M. quadrata 26
subsp. hyperborea 26
subsp. quadrata 26
M. romanica 26
MARCHANTIACEAE 7, 26
MARCHANTIALES 7, 25
JOURNAL OF BRYOLOGY
Marchantioideae 7, 26
Marchantiidae 7
Marchantiophyta 2, 6, 8
Marchantiopsida 7, 25
Marchesinia 23
M. subgenus Marchesinia 23
M. mackaii 23
Marsupella 16
M. adusta = Gymnomitrion adustum 16
M. alpina = Gymnomitrion alpinum 16
M. andreaeoides 16
M. aquatica 16
M. arctica 16
M. boeckii 16
M. brevissima = Gymnomitrion brevissimum 16
M. commutata = Gymnomitrion commutatum 16
M. condensata 16
M. emarginata 16
subsp. aquatica = Marsupella aquatica 16
subsp. tubulosa = Marsupella tubulosa 17
var. aquatica = Marsupella aquatica 16
var. pearsonii = Marsupella emarginata 16
M. funckii 17
var. badensis = Marsupella funckii 17
M. profunda 17
M. ramosa = Marsupella funckii 17
M. sparsifolia 17
[subsp. childii] 17
subsp. sparsifolia 17
M. sphacelata 17
M. spiniloba 17
M. sprucei 17
var. neglecta = Marsupella sprucei 17
var. ustulata = Marsupella sprucei 17
M. stableri 17
M. subemarginata 17
M. tubulosa 17
Massularia = Heterogemma 12
M. incisa = Schistochilopsis incisa 15
M. laxa = Heterogemma laxa 12
Mastigophora 20
M. woodsii 20
MASTIGOPHORACEAE 6, 20
Meesia 48
M. hexasticha 48
M. longiseta 48
M. minor 116
M. minutissima 116
M. triquetra 48
M. uliginosa 48
MEESIACEAE 7, 48
Merceyoideae 36
Mesoptychia 17
M. badensis 17
M. bantriensis 17
subsp. bantriensis 18
subsp. wallfischii 18
M. collaris 18
M. fitzgeraldiae 18
M. gillmanii 18
var. gillmanii 18
var. laxa 5, 18
M. heterocolpos 18
var. arctica 18
var. harpanthoides 18
var. heterocolpos 18
M. rutheana 18
M. sahlbergii 18
M. turbinata 18
Mesoptychioideae 17
Metzgeria 24
M. conjugata 24
subsp. simplex = Metzgeria simplex 24
var. alipila = Metzgeria conjugata 24
var. macvicari = Metzgeria conjugata 24
M. consanguinea 24, 67
M. fruticulosa auct. = Metzgeria violacea 24
M. fruticulosa = Riccardia palmata 24
M. furcata 24
var. expansa = Metzgeria furcata 24
var. flexipilis = Metzgeria furcata 24
[var. pacifica] 24
var. ulvula = Metzgeria furcata 24
[M. hamata = Metzgeria procera] 24
M. hamata auct. = Metzgeria leptoneura 24
M. leptoneura 24
[var. breviseta] 24
var. leptoneura 24
[var. polychaeta] 24
[M. procera] 24
M. pubescens 24
[M. setigera] 24
M. simplex 24
M. temperata auct. = Metzgeria consanguinea 24
[M. temperata] 24, 67
M. violacea 24
METZGERIACEAE 6, 24
METZGERIALES 6, 23
Metzgeriidae 6
Microbryum 38
M. curvicollum 38
M. davallianum 38
var. commutatum 38
var. conicum 38
var. davallianum 38
M. floerkeanum 38
M. fosbergii 38
M. longipes 38
M. rectum 38
M. starckeanum 38
M. vlassovii 38
Microcampylopus 33
M. laevigatus 33
Microeurhynchium 60
M. pumilum 60
Microhypnum 57
M. sauteri 57
Microlejeunea 23
[M. diversiloba] 67
M. ulicina 23
MICROMITRIACEAE 7, 32
Micromitrium 32
M. tenerum 32
Mielichhoferia 51
M. elongata 51
M. mielichhoferiana 51
Mielichhoferioideae 51
MNIACEAE 5, 8, 51
Mnioideae 5, 52
Mnioloma 16
M. subgenus Caracoma 16
M. fuscum 16
Mnium 5, 6, 52
M. blyttii 52
101
102
N. G. HODGETTS ET AL.
M. heterophyllum 52
M. latifolium = Ptychostomum schleicheri var.
latifolium 6
M. hornum 52
M. latifolium = Ptychostomum schleicheri var. latifolium 6
M. lycopodioides 52
M. marginatum 52
var. dioicum 52
var. marginatum 52
M. spinosum 52
M. spinulosum 52
M. stellare 52
M. thomsonii 52
Moerckia 24
M. blyttii 24
M. flotoviana 24
M. hibernica 24
MOERCKIACEAE 7, 24
Molendoa 42
M. hornschuchiana 42
M. schliephackei 42
[M. seravschanica] 69
M. sendtneriana = Molendoa hornschuchiana 42
M. taeniatifolia 42
M. tenuinervis = Molendoa hornschuchiana 42
M. warburgii 42
Mylia 21
M. subgenus Anomalae 21
M. subgenus Mylia 21
M. anomala 21
M. taylorii 21
MYLIACEAE 6, 21
Myliineae 6
Myrinia 56
M. pulvinata 56
MYRINIACEAE 8, 56
Myriocoleopsis 22
M. minutissima 22
subsp. minutissima 22
[subsp. myriocarpa] 22
Myurella 56
M. julacea 56
M. sibirica 56
M. tenerrima 56
Myurium 65
M. hochstetteri 65
MYURIACEAE 8, 65
Myuroclada 61
M. longiramea 61
M. maximowiczii 4, 61
Nardia 17
N. breidleri 17
N. compressa 17
N. geoscyphus 17
var. bifida 17
[var. dioica] 17
var. geoscyphus 17
var. suberecta 17
N. insecta 17
N. japonica 17
N. pacifica 17
N. scalaris 17
[var. botryoidea] 17
[var. harae] 17
var. scalaris 17
Nardioideae 17
Neckera 64
N. baetica = Exsertotheca baetica 64
N. besseri = Alleniella besseri 64
N. cephalonica 64
N. complanata = Alleniella complanata 64
N. crispa = Exsertotheca crispa 64
N. intermedia = Exsertotheca intermedia 64
N. menziesii 64
N. oligocarpa 64
N. pennata 64
N. pumila 64
N. webbiana = Pseudomalia webbiana 65
NECKERACEAE 2, 8, 64, 65
Neoorthocaulis 9
N. attenuatus 9
N. binsteadii 10
N. floerkei 10
N. hyperboreus 10
Niphotrichum = Racomitrium subgenus Niphotrichum 44
N. canescens = Racomitrium canescens 44
subsp. latifolium = Racomitrium canescens subsp.
latifolium 44
N. elongatum = Racomitrium elongatum 44
N. ericoides = Racomitrium ericoides 44
N. panschii = Racomitrium panschii 44
Nobregaea 60
N. latinervis 60
Nogopterium 64
N. gracile 64
NOTOTHYLADACEAE 6, 8
NOTOTHYLADALES 6, 8
Notothylatidae 6
Notothylas 8
N. subgenus Notothylas 8
N. orbicularis 8
Notothyloideae 8
Nowellia 11
N. section Nowellia 11
N. curvifolia 11
Nyholmiella 53
N. gymnostoma 54
N. obtusifolia 54
Obtusifolium 11, 12
O. obtusum 12
Ochyraea = Plathypnum 6, 57
O. tatrensis = Platyhypnum tatrense 6, 58
Octodiceras = Fissidens subgenus Octodiceras 34
O. fontanum = Fissidens fontanus 34
Odontoschisma 11
O. section Cladopodiella 11
O. section Denudata 11
O. section Neesii 11
O. section Odontoschisma 11
O. denudatum 11
subsp. denudatum 11
[subsp. naviculare] 11
[subsp. sandvicense] 11
O. elongatum 11
O. fluitans 11
O. francisci 11
O. macounii 11
O. prostratum = Odontoschisma sphagni 11
O. sphagni 11
Odontoschismatoideae 11
OEDIPODIACEAE 7, 29
OEDIPODIALES 7, 29
Oedipodiella 32
O. australis 32
JOURNAL OF BRYOLOGY
Oedipodiopsida 7, 29
Oedipodium 29
O. griffithianum 29
Oleolophozia 12
O. perssonii 12
Oligotrichum 29
O. hercynicum 29
ONCOPHORACEAE = RHABDOWEISIACEAE 35
Oncophorus 35
O. compactus = Oncophorus wahlenbergii 35
O. demetrii 35
O. dendrophilus 35
O. elongatus 35
O. integerrimus 35
O. riparius = Kiaeria riparia 35
O. virens 35
var. elongatus = Oncophorus integerrimus 35
O. wahlenbergii 35
var. compactus = Oncophorus wahlenbergii 35
Oreas 35
O. martiana 35
Oreoweisia 35
O. torquescens 35
Orthocaulis 10
O. atlanticus 10
O. attenuatus = Neoorthocaulis attenuatus 9
O. binsteadii = Neoorthocaulis binsteadii 10
O. cavifolius 10
O. floerkei = Neoorthocaulis floerkei 10
O. hyperboreus = Neoorthocaulis hyperboreus 10
Orthodicranum = Dicranum 35
O. flagellare = Dicranum flagellare 35
O. fulvum = Dicranum fulvum 35
O. montanum = Dicranum montanum 35
O. scottianum = Dicranum scottianum 35
O. tauricum = Dicranum tauricum 35
ORTHODONTIACEAE 8, 55
ORTHODONTIALES 8, 55
Orthodontium 55
O. gracile 55
O. lineare 4, 55
O. pellucens 55
Orthogrimmia = Grimmia 45
O. alpestris = Grimmia alpestris 45
O. arenaria = Grimmia arenaria 45
O. caespiticia = Grimmia caespiticia 45
O. donniana = Grimmia donniana 45
O. montana = Grimmia montana 45
O. sessitana = Grimmia reflexidens 45
O. triformis = Grimmia triformis 45
Ortholimnobium 56
O. handelii 56
Orthomnion = Plagiomnium 52
O. affine = Plagiomnium affine 52
O. confertidens = Plagiomnium confertidens 52
O. curvatulum = Plagiomnium curvatulum 52
O. cuspidatum = Plagiomnium cuspidatum 52
O. drummondii = Plagiomnium drummondii 52
O. elatum = Plagiomnium elatum 52
O. ellipticum = Plagiomnium ellipticum 52
O. medium = Plagiomnium medium 52
O. undulatum = Plagiomnium undulatum 53
var. madeirense = Plagiomnium undulatum var.
madeirensis 53
O. rostratum = Plagiomnium rostratum 53
Orthothecium 56
O. chryseon 56
O. intricatum 56
O. lapponicum 56
O. rufescens 56
O. strictum 56
ORTHOTRICHACEAE 2, 8, 53
ORTHOTRICHALES 8, 53
Orthotrichoideae 53
Orthotrichum 54
O. acuminatum = Lewinskya acuminata 53
O. affine = Lewinskya affine 53
var. bohemicum = Lewinskya affine 53
O. alpestre 54
O. anomalum 54
O. bistratosum 54
O. callistomum 54
O. cambrense 54
O. casasianum 54
O. columbicum 54
O. comosum 54
[O. consimile] 54
O. consimile auct. eur. = Orthotrichum columbicum 54
O. consobrinum 54
O. crenulatum 54
O. cupulatum 54
var. bistratosum = Orthotrichum bistratosum 54
var. cupulatum 54
var. fuscum 54
var. riparium 54
O. dagestanicum 54
O. dentatum 54
O. diaphanum 54
O. elegans = Lewinskya elegans 53
O. fastigiatum = Lewinskya fastigiata 53
O. gymnostomum = Nyholmiella gymnostoma 54
O. handiense 54
O. hispanicum 54
O. holmenii = Orthotrichum sibiricum 54
O. ibericum = Lewinskya iberica 53
O. iwatsukii = Lewinskya iwatsukii 53
O. laevigatum = Lewinskya laevigata 53
var. japonicum = Lewinskya iwatsukii 53
O. lyellii = Pulvigera lyellii 54
O. macounii 53
subsp. japonicum = Lewinskya iwatsukii 53
O. macrocephalum 54
O. microcarpum 54
O. moravicum 54
O. obtusifolium = Nyholmiella obtusifolia 54
O. pallens 54
O. patens 54
O. pellucidum 54
O. philibertii 54
O. pulchellum 54
O. pumilum 54
O. pylaisii = Lewinskya pylaisii 53
O. rivulare 54
O. rogeri 54
O. rupestre = Lewinskya rupestris 53
O. scanicum 54
O. schimperi 54
O. shawii = Lewinskya shawii 53
O. shevockii 54
O. sibiricum 54
O. sordidum = Lewinskya sordida 53
O. speciosum = Lewinskya speciosa 53
var. brevisetum = Lewinskya breviseta 53
O. sprucei 54
103
104
N. G. HODGETTS ET AL.
O. stellatum 54
O. stramineum 54
O. striatum = Lewinskya striata 53
O. tenellum 54
O. tortidontium = Lewinskya tortidontia 53
[O. urnaceum] 69
O. urnigerum 54
O. vittii 54
O. vladikavkana = Lewinskya vladikavkana 53
Osculatia = Bryum 3, 48
O. apiculata = Anomobryum apiculatum 48
O. barnesii = Bryum dichotomum 48
O. bicolor = Bryum dichotomum 48
O. bornholmensis = Ptychostomum bornholmense 50
O. caespiticia = Ptychostomum imbricatulum 50
O. demaretiana = Bryum demaretianum 48
O. dichotoma = Bryum dichotomum 48
O. gemmifera = Bryum gemmiferum 49
O. gemmilucens = Bryum gemmilucens 49
O. klinggraeffii = Bryum klinggraeffii 49
O. kunzei = Ptychostomum kunzei 51
O. radiculosa = Bryum radiculosum 49
O. rubens = Ptychostomum rubens 51
O. ruderalis = Bryum ruderale 49
O. sauteri = Bryum sauteri 49
O. subapiculata = Imbribryum subapiculatum 49
O. tenuiseta = Imbribryum tenuisetum 49
O. valparaisense = Bryum valparaisense 49
O. violacea = Bryum violaceum 49
Oxymitra 26
O. incrassata 26
OXYMITRACEAE 7, 26
Oxyrrhynchium 60
O. hians 60
O. pumilum = Microeurhynchium pumilum 60
O. schleicheri 60
O. speciosum 60
Oxystegus = Chionoloma 40
O. daldinianus = Chionoloma daldinianum 40
O. hibernicus = Chionoloma hibernicum 40
O. minor = Chionoloma minus 41
O. recurvifolius = Chionoloma recurvifolium 41
O. tenuirostris = Chionoloma tenuirostre 41
var. holtii = Chionoloma tenuirostre var. holtii 41
Palamocladium 59
P. euchloron 59
[P. leskeoides] 69
Pallavicinia 24
P. lyellii 24
PALLAVICINIACEAE 7, 24
PALLAVICINIALES 7, 24
Pallaviciniineae 7
Pallavicinioideae 24
Paludella 48
P. squarrosa 48
Palustriella 56
P. commutata 56
var. sulcata = Palustriella falcata 56
P. decipiens 56
P. falcata 56
P. pluristratosa = Palustriella falcata 56
Paraleptodontium recurvifolium = Chionoloma
recurvifolium 41
Paraleucobryum 35
P. enerve 35
P. longifolium 35
var. sauteri = Paraleucobryum sauteri 35
P. sauteri 35
Pedinophyllum 20
P. interruptum 20
Pelekium 59
P. atlanticum 59
P. minutulum 59
Pellia 25
P. endiviifolia = Apopellia endiviifolia 24
P. epiphylla 25
subsp. borealis 25
subsp. epiphylla 25
P. neesiana 25
PELLIACEAE 7, 24
PELLIALES 7, 24
Pelliidae 6
Peltolepis 25
P. quadrata 25
[Perssonia sanguinea] 69
PETALOPHYLLACEAE 7, 24
Petalophyllum 24
P. ralfsii 24
Phaeoceros 8
P. bulbiculosus = Phymatoceros bulbiculosus 8
P. carolinianus 8
P. laevis 8
subsp. carolinianus = Phaeoceros carolinianus 8
Phaeocerotideae 6, 8
Phascum = Tortula 39
P. cuspidatum = Tortula acaulon 39
var. cuspidatum = Tortula acaulon var. acaulon 39
var. papillosum = Tortula acaulon var. papillosa 39
var. piliferum = Tortula acaulon var. pilifera 39
var. retortifolium = Tortula acaulon var. retortifolia 39
var. schreberianum = Tortula acaulon var.
schreberiana 39
P. vlassovii = Microbryum vlassovii 38
Philonotis 47
P. section Bartramidula 47
P. section Homomorphae 47
P. section Philonotis 47
P. section Philonotula 47
P. arnellii = Philonotis capillaris 47
P. caespitosa 47
P. calcarea 47
P. capillaris 47
P. cernua 47
P. falcata 47
P. fontana 47
P. hastata 47
[P. laxitexta] 69
P. marchica 47
[P. nanothecioidea] 69
P. rigida 47
P. seriata 47
P. tomentella 47
P. uncinata 47
P. yezoana 47
Phymatoceros 8
P. bulbiculosus 8
PHYMATOCEROTACEAE 6, 8
PHYMATOCERATALES 6, 8
Physcomitrella = Physcomitrium 31
P. patens = Physcomitrium patens 31
P. readeri = Physcomitrium readeri 32
Physcomitridium = Physcomitrium 31, 32
P. readeri = Physcomitrium readeri 32
Physcomitrium 31
JOURNAL OF BRYOLOGY
P. arenicola 31
P. eurystomum 31
subsp. acuminatum 31
subsp. eurystomum 31
[P. immersum] 69
[P. niloticum] 69
P. patens 31
P. pyriforme 31
P. readeri 32
P. sphaericum 32
P. × stevensonii 116
PILOTRICHACEAE 8, 55
Plagiobryoides = Ptychostomum 3, 50
P. cellularis = Ptychostomum cellulare 50
Plagiobryum = Ptychostomum 3, 50
P. demissum = Ptychostomum demissum 50
P. zieri = Ptychostomum zieri 51
Plagiochasma 25
P. subgenus Micropylum 25
P. subgenus Plagiochasma 25
P. appendiculatum 25
[P. eximium] 69
[P. microcephalum var. tunesicum] 69
P. rupestre 25
var. rupestre 25
[var. volkii] 25
Plagiochila 20
P. section Arrectae 20
P. section Fuscoluteae 20
P. section Glauscentes 20
P. section Plagiochila 20
P. section Poeltiae 21
P. section Rutilantes 21
P. section Vagae 21
[P. ambagiosa = Plagiochila spinulosa] 67
P. arctica 20
subsp. arctica = Plagiochila arctica 20
var. arctica 20
[var. intermedia] 20
P. asplenioides 21
subsp. arctica = Plagiochila arctica 20
subsp. porelloides = Plagiochila porelloides 21
var. devexa = Plagiochila porelloides var. porelloides 21
var. humilis = Plagiochila porelloides var. porelloides 21
var. lobata = Plagiochila asplenioides var. arctica 20
var. minor = Plagiochila porelloides var. porelloides 21
var. subarctica = Plagiochila porelloides var. subarctica 21
P. atlantica = Plagiochila heterophylla var. heterophylla 20
P. bifaria 20
var. bifaria 20
[var. rosea] 20
P. britannica 21
P. carringtonii 21
subsp. carringtonii 21
[subsp. lobuchensis] 21
P. dubia auct. eur. = P. virginica 21
[P. dubia = Plagiochila patula] 67
P. exigua 21
P. heterophylla 20
[var. beauverdii] 20
var. heterophylla 20
[P. javanica] 67
P. longispina 20
P. maderensis 21
P. major = Plagiochila asplenioides 21
P. norvegica = Plagiochila porelloides var. norvegica 21
P. papillifolia 20
[P. patula] 21, 67
P. porelloides 21
var. norvegica 21
var. porelloides 21
var. subarctica 21
P. punctata 20
P. retrorsa 20
P. spinulosa 20
P. stricta 20
[P. uniformis] 67
P. virginica 21, 67
[var. caroliniana] 21
[var. euryphylla] 21
var. virginica 21
PLAGIOCHILACEAE 6, 20
Plagiomnium 52
P. section Plagiomnium 52
P. section Rostrata 53
P. section Undulata 52
P. affine 52
P. confertidens 52
P. curvatulum 52
P. cuspidatum 52
P. drummondii 52
P. elatum 52
P. ellipticum 52
P. medium 52
P. rostratum 53
P. undulatum 53
var. madeirense 53
var. undulatum 53
Plagiopus 47
P. oederianus 47
var. alpinus 47
var. oederianus 47
PLAGIOTHECIACEAE 8, 55
Plagiothecium 55
P. berggrenianum 55
P. cavifolium 55
P. curvifolium 55
P. denticulatum 55
var. denticulatum 55
var. obtusifolium 55
var. undulatum 55
P. handelii = Ortholimnobium handelii 56
P. laetum 55
P. latebricola 55
P. neckeroideum 56
P. nemorale 56
P. noricum = Plagiothecium neckeroideum 56
P. piliferum 56
P. platyphyllum 56
P. rossicum 56
P. succulentum 56
P. svalbardense 56
P. undulatum 56
Plasteurhynchium 59
P. meridionale 59
P. striatulum 59
Platydictya 56
P. jungermannioides 56
[Platygyrella densa] 69
Platygyrium 62
P. repens 62
Platyhypnidium = Rhynchostegium 59
P. grolleanum = Rhynchostegium riparioides 60
P. lusitanicum = Rhynchostegium alopecuroides 59
105
106
N. G. HODGETTS ET AL.
P. mutatum = Rhynchostegium alopecuroides 59
P. torrenticola = Rhynchostegium riparioides 60
Platyhypnum 6, 57
P. alpestre 57
P. alpinum 57
P. cochlearifolium 58
P. duriusculum 58
P. molle 58
P. montanum = Campylophyllum montanum 57
P. norvegicum 58
P. smithii 58
P. tatrense 6, 58
[Plaubelia sprengelii] 70
Plectocolea = Solenostoma 18
P. hyalina = Solenostoma hyalinum 18
P. obovata = Solenostoma obovatum 18
P. subelliptica = Solenostoma subellipticum 18
Plenogemma 54
P. phyllantha 54
Pleuridium 36
P. acuminatum 36
P. palustre = Cleistocarpidium palustre 36
P. subulatum 36
Pleurochaete = Tortella 42
P. squarrosa = Tortella squarrosa 42
Pleurocladula = Fuscocephaloziopsis 10
P. affinis = Fuscocephaloziopsis affinis 10
P. albescens = Fuscocephaloziopsis albescens 10
var. islandica = Fuscocephaloziopsis albescens var.
islandica 10
P. catenulata = Fuscocephaloziopsis catenulata 10
P. connivens = Fuscocephaloziopsis connivens 10
P. leucantha = Fuscocephaloziopsis leucantha 11
P. loitlesbergeri = Fuscocephaloziopsis loitlesbergeri 11
P. lunulifolia = Fuscocephaloziopsis lunulifolia 11
P. macrostachya = Fuscocephaloziopsis macrostachya 11
P. pleniceps = Fuscocephaloziopsis pleniceps 11
Pleurozia 24
P. subgenus Constantifolia 24
[P. gigantea] 69
P. purpurea 24
PLEUROZIACEAE 6, 24
PLEUROZIALES 6, 24
Pleurozium 64
P. schreberi 64
Pogonatum 29
P. aloides 29
P. dentatum 29
P. nanum 29
P. neesii 29
P. urnigerum 29
Pohlia 52
[P. alba] 70
P. andalusica 52
P. andrewsii 52
P. annotina 52
P. atropurpurea 52
P. beringiensis 52
P. bolanderi 52
P. bulbifera 52
P. camptotrachela 52
P. cruda 52
P. crudoides 52
P. drummondii 52
P. elongata 52
var. acuminata 52
var. elongata 52
var. greenii 52
P. erecta 52
P. filum 52
P. flexuosa 52
var. flexuosa 52
var. pseudomuyldermansii 52
P. inclinata = Ptychostomum inclinatum 50
P. lescuriana 52
P. longicolla 52, 68
P. ludwigii 52
P. lutescens 52
P. melanodon 52
P. nutans 52
subsp. nutans 52
subsp. schimperi 52
P. obtusifolia 52
P. proligera 52
[P. philonotula] 68
[P. saprophila] 68, 70
P. scotica 52
P. sphagnicola 52
P. tundrae 52
P. vexans 52
P. wahlenbergii 52
var. calcarea 52
var. glacialis 52
var. wahlenbergii 52
POLYTRICHACEAE 2, 7, 29
POLYTRICHALES 7, 29
Polytrichastrum 29
P. alpinum 29
var. fragile = Polytrichastrum fragile 30
var. septentrionale = Polytrichastrum septentrionale 30
P. altaicum 29
P. formosum = Polytrichum formosum 30
var. densifolium = Polytrichum densifolium 30
P. fragile 30
P. longisetum = Polytrichum longisetum 30
P. norwegicum = Polytrichastrum alpinum 29
P. pallidisetum = Polytrichum pallidisetum 30
P. septentrionale 30
P. sexangulare 30
var. vulcanicum = Polytrichastrum sphaerothecium 30
P. sphaerothecium 30
Polytrichopsida 7, 29
Polytrichum 30
P. alpestre = Polytrichum strictum 30
P. alpinum = Polytrichastrum alpinum 29
var. arcticum = Polytrichastrum alpinum 29
[P. appressum = Polytrichum subpilosum] 30
P. commune 30
var. commune = Polytrichum commune 30
var. humile = Polytrichum perigoniale 30
var. perigoniale = Polytrichum perigoniale 30
var. swartzii = Polytrichum swartzii 30
var. uliginosum = Polytrichum commune 30
P. densifolium 30
P. formosum 30
var. densifolium = Polytrichum densifolium 30
P. fragile = Polytrichastrum fragile 30
P. hyperboreum 30
P. jensenii 30
P. juniperinum 30
P. longisetum 30
P. pallidisetum 30
P. perigoniale 30
P. piliferum 30
JOURNAL OF BRYOLOGY
P. septentrionale = Polytrichastrum septentrionale 30
P. sexangulare = Polytrichastrum sexangulare 30
P. sphaerothecium = Polytrichastrum sphaerothecium 30
P. strictum 30
[P. subpilosum] 30
P. swartzii 30
P. uliginosum = Polytrichum commune 30
Porella 23
P. arboris-vitae 23
subsp. arboris-vitae 23
[subsp. nitidula] 23
var. killarniensis = Porella arboris-vitae subsp.
arboris-vitae 23
var. obscura = Porella arboris-vitae subsp. arboris-vitae 23
P. baueri 23
P. canariensis 23
P. cordaeana 23
var. faeroensis = Porella cordaeana 23
var. simplicior = Porella cordaeana 23
P. inaequalis 23
[P. navicularis] 67
P. obtusata 23
P. pinnata 23
P. platyphylla 23, 67
var. subsquarrosa = Porella platyphylla 23
[P. platyphylloidea] 67
PORELLACEAE 6, 23
PORELLALES 6, 21
Porellineae 6
Pottia = Tortula 39
P. commutata = Microbryum davallianum var.
commutatum 38
P. conica = Microbryum davallianum var. conicum 38
P. crinita = Tortula viridifolia 40
P. intermedia = Tortula caucasica 39
P. lanceolata = Tortula lindbergii 40
P. starckeana = Microbryum starckeanum 38
subsp. conica = Microbryum davallianum var. conicum 38
POTTIACEAE 2, 7, 36
Pottioideae 36
Pottiopsis 42
P. caespitosa 42
Prasanthus 17
[P. jamalicus] 69
P. suecicus 17
Preissia = Marchantia subgenus Preissia 26
P. quadrata = Marchantia quadrata 26
subsp. hyperborea = Marchantia quadrata subsp.
hyperborea 26
Protobryum = Tortula 39
Protobryum bryoides = Tortula protobryoides 40
Protolophozia 12
P. elongata 12
P. herzogiana 12
Pseudanomodon 64
P. attenuatus 64
Pseudephemerum 36
P. nitidum 36
Pseudoamblystegium 58
P. subtile 58
Pseudobryum 53
P. cinclidioides 53
Pseudocalliergon = Drepanocladus 57
P. angustifolium = Drepanocladus angustifolius 57
P. brevifolium = Drepanocladus brevifolius 57
P. lycopodioides = Drepanocladus lycopodioides 57
P. trifarium = Drepanocladus trifarius 57
P. turgescens = Drepanocladus turgescens 57
Pseudocampylium 58
P. radicale 58
Pseudocrossidium 38
P. hornschuchianum 38
P. obtusulum 38
P. replicatum 38
P. revolutum 38
Pseudohygrohypnum 63
P. eugyrium 63
P. fertile 63
P. subeugyrium 63
Pseudoleskea = Lescuraea 59
P. artariae = Pseudoleskeopsis artariae 59
P. incurvata = Lescuraea incurvata 59
P. patens = Lescuraea patens 59
P. radicosa = Lescuraea radicosa 59
P. saviana = Lescuraea saviana 59
PSEUDOLESKEACEAE 8, 59
Pseudoleskeella 59
P. catenulata 59
P. nervosa 59
P. papillosa 59
P. rupestris 59
P. tectorum 59
PSEUDOLESKEELLACEAE 8, 59
Pseudoleskeopsis 59
P. artariae 59
[P. bollei] 70
[P. pseudoattenuata] 70
Pseudolophozia = Barbilophozia subgenus Sudeticae 9
P. debiliformis = Barbilophozia sudetica 9
P. sudetica = Barbilophozia sudetica 9
Pseudomalia 65
P. webbiana 65
Pseudomarsupidium 9
P. decipiens 9
Pseudorhynchostegiella 60
P. duriaei 60
Pseudoscleropodium 59
P. purum 59
Pseudostereodon 63
P. procerrimus 63
[Pseudosymblepharis] 40
P. bombayensis = Chionoloma bombayense 40
Pseudotaxiphyllum 56
P. elegans 56
P. laetevirens 56
Pseudotritomaria 14
P. heterophylla 14
Psilopilum 30
P. cavifolium 30
P. laevigatum 30
PTERIGYNANDRACEAE 8, 56
Pterigynandrum 56
P. filiforme 56
var. majus = Pterigynandrum filiforme 56
Pterogonium = Nogopterium 64
P. gracile = Nogopterium gracile 64
Pterygoneurum 39
P. compactum = Pterygoneurum ovatum 39
P. crossidioides = Pterygoneurum ovatum 39
P. kozlovii 39
P. lamellatum 39
P. ovatum 39
P. papillosum 39
P. sampaianum 39
107
108
N. G. HODGETTS ET AL.
P. squamosum = Pterygoneurum ovatum 39
P. subsessile 39
PTILIDIACEAE 6, 23
PTILIDIALES 6, 23
Ptilidium 23
P. ciliare 23
P. pulcherrimum 23
var. subpinnatum = Ptilidium pulcherrimum 23
Ptilium 63
P. crista-castrensis 63
Ptychanthoideae 23
Ptychodium = Lescuraea 59
P. plicatum = Lescuraea plicata 59
PTYCHOMITRIACEAE 7, 43
Ptychomitrium 44
P. incurvum 44
P. nigrescens 44
P. polyphyllum 44
Ptychostomum 3, 6, 50
P. archangelicum = Ptychostomum inclinatum 50
P. arcticum 50
P. austriacum 6, 50
P. boreale = Ptychostomum pallescens 51
P. bornholmense 50
P. calophyllum 50
P. capillare 50
P. cellulare 6, 50
P. cernuum 50
P. compactum 50
var. compactum 50
var. rutheanum 50
P. creberrimum 50
P. cryophilum 50
P. cyclophyllum 50
P. demissum 50
P. donianum 50
P. elegans 6, 50
P. funkii 50
P. imbricatulum 3, 50
P. imbricatum = Ptychostomum inclinatum 50
P. inclinatum 50
P. intermedium 50
P. knowltonii 50
P. kunzei 50
P. longisetum 51
P. minii 6, 51
P. moravicum 51
P. neodamense = Ptychostomum pseudotriquetrum 51
P. pallens 51
P. pallescens 51
P. pseudotriquetrum 3, 51
var. bimum 51
var. pseudotriquetrum 51
P. rubens 51
P. salinum 51
P. schleicheri 6, 51
var. latifolium 6, 51
var. schleicheri 51
P. torquescens 51
P. turbinatum 51
P. warneum 51
P. weigelii 51
P. wrightii 51
P. zieri 51
Pulvigera 54
P. lyellii 54
Pylaisia 63
P. polyantha 63
P. selwynii 63
PYLAISIACEAE 8, 63
PYLAISIADELPHACEAE 8, 62
Pyramidula 31
P. tetragona 31
Pyramiduloideae 31
Racomitrioideae 44
Racomitrium 44
R. subgenus Cataracta 44
R. subgenus Ellipticodryptodon 44
R. subgenus Niphotrichum 44
R. subgenus Racomitrium 44
R. aciculare 44
R. affine 44
R. aquaticum 44
R. canescens 44
subsp. canescens 44
subsp. latifolium 44
R. ellipticum 44
R. elongatum 44
R. ericoides 44
R. fasciculare 44
R. hespericum 44
R. heterostichum 44
R. himalayanum 44
R. lamprocarpum 44
R. lanuginosum 44
R. lusitanicum 44
R. macounii 44
subsp. alpinum 44
subsp. macounii 44
R. microcarpon 44
R. nivale 44
R. obtusum 44
R. panschii 44
R. sudeticum 44
Radula 23
R. subgenus Radula 23
R. subgenus Volutoradula 23
R. aquilegia 23
R. carringtonii 23
R. complanata 23
subsp. lindenbergiana = Radula lindenbergiana 23
var. alpestris = Radula complanata 23
R. holtii 23
R. jonesii 23
R. lindbergiana = Radula lindenbergiana 23
R. lindenbergiana 23
R. nudicaulis 23
var. delicatula 23
var. nudicaulis 23
[R. prolifera] 69
R. voluta 23
R. visianica 23
R. wichurae 23
RADULACEAE 6, 23
Radulineae 6
Reboulia 25
R. hemisphaerica 25
[subsp. acrogyna] 25
subsp. australis 25
subsp. dioica 25
subsp. hemisphaerica 25
[subsp. orientalis] 25
[subsp. paradoxa] 25
[var. fissisquama] 25
JOURNAL OF BRYOLOGY
[var. turkestanica] 25
Rhabdoweisia 35
R. crenulata 35
R. crispata 35
R. fugax 35
RHABDOWEISIACEAE; 7, 35
Rhamphidium 36
R. purpuratum 36
RHIZOGONIACEAE 8, 55
RHIZOGONIALES 8, 55
Rhizomnium 53
R. andrewsianum 53
R. gracile 53
R. magnifolium 53
R. pseudopunctatum 53
R. punctatum 53
var. hermanperssonii 53
var. punctatum 53
[R. striatulum] 70
Rhodobryum 3, 51
R. ontariense 51
R. roseum 51
Rhynchostegiella 60
R. azorica 60
R. bourgaeana 60
R. curviseta 60
R. durieui = Pseudorhynchostegiella durieui 60
R. jacquinii = Rhynchostegiella teneriffae 60
R. litorea 60
R. macilenta = Rhynchostegiella teneriffae 60
R. pseudolitorea 60
R. teesdalei = Rhynchostegiella teneriffae 60
R. tenella 60
var. meridionalis = Rhynchostegiella litorea 60
R. teneriffae 60
R. tenuicaulis = Brachythecium tenuicaule 60
R. trichophylla 60
R. tubulosa 60
Rhynchostegium 59
R. alopecuroides 59
R. arcticum = Rhynchostegium murale 60
R. confertum 59
R. confusum 59
R. megapolitanum 60
R. murale 60
R. riparioides 60
R. rotundifolium 60
R. strongylense 60
RHYTIDIACEAE 8, 64
Rhytidiadelphus 64
R. loreus 64
R. squarrosus 64
R. subpinnatus 64
R. triquetrus = Hylocomiadelphus triquetrus 63
Rhytidium 64
R. rugosum 64
Riccardia 23
R. subgenus Riccardia 23
R. section Riccardia 23
R. chamedryfolia 23
var. major = Riccardia chamedryfolia 23
var. submersa = Riccardia chamedryfolia 23
R. incurvata 23
R. latifrons 23
subsp. arctica 23
subsp. latifrons 23
[var. miyakeana] 23
109
R. multifida 23
[subsp. decrescens] 23
subsp. multifida 23
[subsp. synoica] 23
R. palmata 23
Riccia 4, 26
R. subgenus Riccia 26
R. subgenus Ricciella 27
R. section Pilifer 27
R. section Riccia 26
R. section Ricciella 27
R. section Spongodes 27
R. atlantica 26
R. atromarginata 26
var. atromarginata 26
[var. jovet-astiae] 26
R. beyrichiana 26
R. bicarinata 26
R. bifurca 26
var. subinermis = Riccia bifurca 26
R. boumanii 27
R. breidleri 26
[R. bullosa] 67
[R. capensis = Riccia concava] 67
R. cavernosa 27
var. angustior = Riccia cavernosa 27
R. canaliculata 27
R. canescens = Riccia ciliata 26
[R. chudoana] 69
R. ciliata 26
var. epilosa = Riccia ciliata 26
var. intumescens = Riccia ciliata 26
var. violacea = Riccia ciliata 26
R. ciliifera 26
[R. concava] 67
[R. congoana] 69
[R. crinita] 26
R. crinita auct. eur. = Riccia ciliata 26
R. crozalsii 26
R. crustata 27
R. crystallina 26
R. dalslandica = Riccia ciliata 26
R. duplex 27
var. duplex 27
[var. megaspora] 27
R. fluitans 27, 68
R. frostii 27
[var. crystallinoides] 27
var. frostii 27
R. glauca 26
var. ciliaris 26
var. glauca 26
var. major = Riccia glauca 26
var. subinermis = Riccia glauca var. ciliaris 26
R. gothica 26
R. gougetiana 26
var. armatissima 26
var. gougetiana 26
R. huebeneriana 27
subsp. huebeneriana 27
[subsp. sullivantii] 27
var. pseudo-frostii = Riccia huebeneriana subsp. huebeneriana 27
R. intumescens = Riccia ciliata 26
R. lamellosa 26
R. ligula 26
[R. limbata] 67
110
N. G. HODGETTS ET AL.
R. lindenbergiana = Riccia sorocarpa subsp. arctica 26
R. macrocarpa 26
[R. mamillata] 69
R. marginata = Riccia beyrichiana 26
R. melitensis = Riccia ciliifera 26
R. michelii 26
R. nigrella 26, 67
R. oelandica = Riccia subbifurca 27
[R. oerstediana] 68
R. papillosa 26
R. perennis 27
[R. polycarpa] 69
R. rhenana 27
var. rhenana 27
var. violacea 27
R. sommieri 26
R. sorocarpa 26
subsp. arctica 4, 26
subsp. erythrophora 5, 27
subsp. sorocarpa 27
var. arctica = Riccia sorocarpa subsp. arctica 4
var. heegii = Riccia sorocarpa 26
R. subbifurca 27
R. teneriffae = Riccia cavernosa 27
R. trabutiana 27
R. trichocarpa = Riccia ciliata 26
R. warnstorfii 27
var. ciliaris= Riccia warnstorfii 27
var. commutata = Riccia warnstorfii 27
var. subinermis = Riccia warnstorfii 27
RICCIACEAE 7, 26
Ricciocarpos 27
R. natans 27
Riella 27
R. subgenus Riella 27
R. subgenus Trabutiella 27
R. affinis 27
R. battandieri 27
R. bialata 27
R. cossoniana 27
[R. cyrenaica] 27, 69
R. echinata 27
R. gallica 27
R. helicophylla 27
var. helicophylla 27
var. macrocarpa 27
R. mediterranea 27
R. notarisii 27
[R. numidica] 69
[R. parisii] 68, 69
R. reuteri 27
[R. sersuensis] 27, 69
RIELLACEAE 7, 27
Roaldia 63
R. dolomitica 63
R. revoluta 63
Rosulabryum = Ptychostomum 3, 50
[R. albolimbatum] 50
[R. billardierei] 50
R. bornholmense = Ptychostomum bornholmense 50
R. canariense = Bryum canariense 48
R. capillare = Ptychostomum capillare 50
R. donianum = Ptychostomum donianum 50
R. elegans = Ptychostomum elegans 50
R. laevifilum = Ptychostomum moravicum 51
R. rubens = Ptychostomum rubens 51
R. torquescens = Ptychostomum torquescens 51
Saccobasis 14
S. polita 14
S. polymorpha 14
Saccogyna 18
S. viticulosa 18
SACCOGYNACEAE 6, 18
Saelania 43
S. glaucescens 43
SAELANIACEAE 7, 43
Sanionia 58
S. nivalis 58, 68
[S. georgicouncinata] 58, 68
S. georgicouncinata auct. eur. = Sanionia nivalis 58
S. orthothecioides 58
S. uncinata 58
Sarmentypnum 58
S. exannulatum 58
S. procerum 58
S. sarmentosum 58
S. trichophyllum 58
S. tundrae 58
Sauteria 25
S. section Sauteria 25
S. alpina 25
Scapania 5, 14
S. subgenus Gracilidae 14
S. subgenus Plicaticalyx 14
S. subgenus Scapania 14
S. section Aequilobae 14
S. section Apiculatae 14
S. section Compactae 14
S. section Curtae 15
S. section Cuspiduligerae 15
S. section Hyperboreae 15
S. section Kaalaasia 15
S. section Nemorosae 15
S. section Planifoliae 14
S. section Scapania 15
S. section Scapaniella 15
S. section Simmonsiae 15
S. section Sphaeriferae 15
S. section Verrucosae 15
S. aequiloba 14
S. apiculata 14
S. aspera 14
S. brevicaulis auct. eur. = Scapania degenii 15
[S. brevicaulis] 15, 69
var. dubia = Scapania degenii 15
S. calcicola 15
S. carinthiaca 14
var. carinthiaca 14
var. massalongi 14
S. compacta 14
S. crassiretis 15
S. curta 15
var. curta 15
var. grandiretis 15
var. isoloba 15
S. cuspiduligera 15
var. cuspiduligera 15
[var. diplophyllopsis] 15
S. degenii 15
var. dubia = Scapania degenii 15
S. glaucocephala 15
var. glaucocephala 14
[var. saxicola] 15
var. scapanoides = Scapania scapanioides 15
JOURNAL OF BRYOLOGY
S.
S.
S.
S.
S.
gracilis 14
gymnostomophila 15
helvetica 15
hyperborea 15
irrigua 15
subsp. irrigua 15
subsp. rufescens 15
var. rubescens = Scapania irrigua subsp. irrigua 15
S. kaurinii 14
S. ligulifolia 15
S. lingulata 15
var. lingulata 15
var. microphylla 15
[S. microdonta] 69
S. mucronata 15
subsp. praetervisa = Scapania praetervisa 15
S. nemorea 15
S. nimbosa 14
S. obcordata 15
S. obscura 15
S. ornithopodioides 14
S. paludicola 5, 15
var. paludicola 15
var. rotundiloba 5, 15
var. rufescens = Scapania paludicola 15
[var. viridigemma] 15
S. paludosa 15
var. isoloba = Scapania paludosa 15
var. rubiginosa = Scapania paludosa 15
var. vogesiaca = Scapania paludosa 15
S. paradoxa = Scapania obcordata 15
S. parvifolia 15
S. praetervisa 15
S. scandica 15
var. argutedentata 15
[var. dimorpha] 15
var. grandiretis 15
var. scandica 15
S. scapanioides 15
S. simmonsii 15
S. sphaerifera 15
S. spitsbergensis 15
S. subalpina 15
[var. haynesiae] 15
[var. muddiae] 15
var. subalpina 15
S. tundrae 15
S. uliginosa 15
S. umbrosa 15
S. undulata 15
var. aequatiformis = Scapania undulata 15
var. dentata = Scapania undulata 15
var. oakesii = Scapania undulata 15
S. verrucosa 15
S. zemliae 15
SCAPANIACEAE 6, 14
Schistidium 46
S. abrupticostatum 46
S. agassizii 46
S. andreaeopsis 46
S. apocarpum 46
S. atrichum 46
S. atrofuscum 46
S. boreale 46
S. brunnescens 46
subsp. brunnescens 46
subsp. griseum 46
S. bryhnii 46
S. canadense 46
[S. cinclidodonteum] 70
S. confertum 46
S. confusum 46
S. convergens 46
S. crassipilum 46
S. crenatum 46
S. dupretii 46
S. echinatum 46
S. elegantulum 46
subsp. elegantulum 46
subsp. wilsonii 46
S. flaccidum 46
S. flexipile 46
S. frigidum 46
var. frigidum 46
var. havaasii 46
S. frisvollianum 46
S. grande 46
S. grandirete 46
S. helveticum 46
S. holmenianum 46
S. lancifolium 46
S. marginale 46
S. maritimum 46
subsp. maritimum 46
subsp. piliferum 46
S. memnonium 116
S. obscurum 46
S. occidentale 46
S. papillosum 46
S. platyphyllum 46
var. abrupticostatum = Schistidium abrupticostatum 46
S. poeltii 46
S. pruinosum 46
S. pulchrum 46
S. recurvum 46
S. rivulare 46
S. robustum 46
S. scandicum 46
S. sibiricum 46
S. sinensiapocarpum 46
S. singarense = Schistidium helveticum 46
S. sordidum 46
S. spinosum 46
S. strictum 46
S. subconfertum 46
S. subflaccidum 46
S. subjulaceum 46
S. submuticum 46
subsp. arcticum 46
subsp. submuticum 46
S. succulentum 46
S. tenerum 46
S. tenuinerve 46
S. trichodon 46
var. nutans 46
var. trichodon 46
S. umbrosum 46
S. venetum 46
Schistochilopsis 15
S. grandiretis 15
S. elegans = Lophoziopsis excisa var. elegans 13
S. hyperarctica 15
S. incisa 15
var. inermis = Schistochilopsis incisa 15
111
112
N. G. HODGETTS ET AL.
S. opacifolia 15
Schistostega 36
S. pennata 36
SCHISTOSTEGACEAE 7, 36
Schizophyllopsis 10
S. sphenoloboides 10
Schizymenium 52
S. pontevedrense 52
Schljakovia 10
S. kunzeana 10
Schljakovianthus 10
S. quadrilobus 10
Sciuro-hypnum 61
S. curtum 61
S. dovrense 61
S. flotowianum 61
S. glaciale 61
S. latifolium 61
S. oedipodium 62
S. ornellanum 62
S. plumosum 62
S. populeum 62
S. reflexum 62
S. starkei 62
S. tromsoeense 62
Scleropodium 62
S. cespitans 62
S. touretii 62
Scopelophila 36
S. cataractae 36
S. ligulata 36
SCORPIDIACEAE 8, 58
Scorpidium 58
S. cossonii 58
S. revolvens 58
S. scorpioides 58
Scorpiurium 60
S. circinatum 60
S. deflexifolium 60
S. sendtneri 60
Seligeria 43
S. subgenus Blindiadelphus = Blindiadelphus 43
S. subgenus Megalosporia 43
S. subgenus Seligeria 43
S. austriaca 43
S. acutifolia 43
S. alpestris = Seligeria patula 43
S. brevifolia 43
S. calcarea 43
S. calycina 43
S. campylopoda = Blindiadelphus campylopodus 43
S. carniolica 43
S. diversifolia = Blindiadelphus diversifolius 43
S. donniana 43
S. galinae = Seligeria donniana 43
S. irrigata 43
S. oelandica 43
S. patula 43
var. alpestris = Seligeria patula 43
S. paucifolia auct. eur. = Seligeria calycina 43
S. polaris = Blindiadelphus polaris 43
S. pusilla 43
S. recurvata = Blindiadelphus recurvatus 43
S. subimmersa = Blindiadelphus subimmersus 43
S. transylvanica = Brachydontium trichodes 43
S. trifaria 43
var. longifolia 43
var. trifaria 43
S. tristichoides 43
var. patula = Seligeria patula 43
SELIGERIACEAE 7, 43
SEMATOPHYLLACEAE 8, 63
Sematophyllum 63
S. adnatum 4, 63
S. demissum 63
S. substrumulosum 63
Serpoleskea 58
S. confervoides 58
Solenostoma 18
S. subgenus Eucalyx 18
S. subgenus Metasolenostoma 18
S. subgenus Plectocolea 18
S. subgenus Solenostoma 18
S. callithrix 18
S. caucasicum 18
S. confertissimum 18
S. gracillimum 18
S. handelii 18
S. hyalinum 18
[S. lignicola] 69
S. obovatum 18
S. paroicum 18
S. pusillum = Solenostoma sphaerocarpum 18
S. sphaerocarpum 18
var. nanum = Solenostoma sphaerocarpum 18
S. subellipticum 18
[S. subtilissimum] 69
SOLENOSTOMATACEAE 6, 18
Southbya 19
S. nigrella 19
S. tophacea 19
SOUTHBYACEAE 6, 19
SPHAEROCARPACEAE 7, 27
SPHAEROCARPALES 7, 27
Sphaerocarpos 27
S. subgenus Austrosphaerocarpos 27
S. subgenus Sphaerocarpos 27
S. europaeus 27, 67
S. michelii 27
S. stipitatus 4, 27
S. texanus auct. = Sphaerocarpos europaeus 27, 68
[S. texanus] 27, 68
SPHAGNACEAE 2, 7, 27
SPHAGNALES 7, 27
SPHAGNOPHYTINA 7
SPHAGNOPSIDA 7, 27
Sphagnum 27
S. subgenus Acutifolia 28
S. subgenus Cuspidata 28
S. subgenus Rigida 28
S. subgenus Sphagnum 28
S. subgenus Subsecunda 28
S. section Acutifolia 28
S. section Insulosa 28
S. section Polyclada 28
S. section Squarrosa 28
S. affine 28
var. flagellare = Sphagnum affine 28
S. angermanicum 28
S. angustifolium 28
S. annulatum 28
S. aongstroemii 28
S. arcticum 28
S. auriculatum 28, 68
JOURNAL OF BRYOLOGY
S. austinii 28
S. balticum 28
S. beothuk 28
S. brevifolium = Sphagnum fallax var. brevifolium 28
S. capillifolium 28
subsp. capillifolium = Sphagnum capillifolium 28
subsp. rubellum = Sphagnum rubellum 28
S. centrale 28
S. compactum 28
S. concinnum 28
S. contortum 28
S. cuspidatum 28
var. cuspidatum 28
var. viride 28
S. denticulatum = Sphagnum auriculatum 28
S. divinum 28
S. fallax 28
subsp. isoviitae = Sphagnum fallax var. isoviitae 28
var. brevifolium 28
var. fallax 28
var. isoviitae 28
S. fimbriatum 28
subsp. concinnum = Sphagnum concinnum 28
S. flexuosum 28
S. fuscum 28
S. girgensohnii 28
[S. imbricatum] 28
subsp. affine = Sphagnum affine 28
subsp. austinii = Sphagnum austinii 28
S. inundatum 28, 68
S. isoviitae = Sphagnum fallax var. isoviitae 28
S. jensenii 29
S. lenense 29
[S. lescurii] 68, 70
S. lindbergii 29
[S. magellanicum Brid.] 28, 68
S. magellanicum auct. eur. p.p. = Sphagnum divinum 28
S. magellanicum auct. eur. p.p. = Sphagnum medium 28
S. majus 29
subsp. majus 29
subsp. norvegicum 29
S. medium 28
S. mirum 28
S. molle 28
S. nitidulum 28
S. obtusum 29
S. olafii 28
S. palustre 28
var. centrale = Sphagnum centrale 28
S. papillosum 28
[S. perfoliatum] 70
S. platyphyllum 28
S. pulchrum 29
S. pylaesii 28
S. quinquefarium 28
S. recurvum 29
S. riparium 29
S. rubellum 28
S. rubiginosum 28
S. russowii 28
S. skyense 28
S. squarrosum 28
S. strictum 28
S. subfulvum 28
subsp. purpureum 28
subsp. subfulvum 28
S. subnitens 28
113
subsp. ferrugineum 28
subsp. subnitens 28
var. ferrugineum = Sphagnum subnitens subsp. ferrugineum 28
S. subsecundum 28
S. tenellum 29
S. teres 28
S. tescorum 28
S. troendelagicum 29
S. tundrae 28
S. venustum 28
S. viride = Sphagnum cuspidatum var. viride 28
S. warnstorfii 28
S. wulfianum 28
Sphenolobopsis 10
S. pearsonii 10
Sphenolobus 10
S. cavifolius = Orthocaulis cavifolius 10
S. minutus 10
S. saxicola 10
SPLACHNACEAE 7, 47
SPLACHNALES 7, 47
Splachnobryum 42
[S. aquaticum] 70
[S. limbatum] 70
S. obtusum 42
Splachnoideae 48
Splachnum 48
S. ampullaceum 48
S. luteum 48
S. melanocaulon 48
S. pensylvanicum 48
S. rubrum 48
S. sphaericum 48
S. vasculosum 48
Stegonia 39
S. latifolia 39
var. latifolia 39
var. pilifera 39
Stereodon 62
S. aemulans 63
S. callichrous 63
S. fertilis = Pseudohygrohypnum fertile 63
S. hamulosus 63
S. holmenii 63
S. pratensis 63
S. subimponens 63
STEREODONTACEAE 8, 62
Straminergon 58
S. stramineum 58
Streblotrichoideae 40
Streblotrichum 40
S. convolutum 40
var. commutatum 40
var. convolutum 40
S. commutatum = Streblotrichum convolutum var.
commutatum 40
S. enderesii 40
Streptocolea = Grimmia 45
S. atrata = Grimmia atrata 45
Syntrichia 39
[S. amphidiacea] 70
S. amplexa = Tortula amplexa 39
S. bogotensis 39
S. calcicola 39
S. caninervis 39
var. abranchesii 39
var. astrakhanica 39
114
N. G. HODGETTS ET AL.
var. caninervis 39
var. gypsophila 39
[var. pseudodesertorum] 70
S. densa = Syntrichia ruralis 39
S. echinata 39
S. fragilis 39
S. glabra = Syntrichia ruralis 39
S. handelii 39
S. laevipila 39
S. latifolia 39
S. minor 39
S. montana 39
var. calva 39
var. montana 39
S. norvegica 39
S. pagorum = Syntrichia laevipila 39
S. papillosa 39
S. papillosissima 39
S. princeps 39
S. rigescens 39
S. ruraliformis 39
S. ruralis 39
var. epilosa 39
var. ruraliformis = S. ruraliformis 39
var. ruralis 39
S. sinensis 39
S. submontana 39
S. subpapillosissima 39
S. virescens 39
Syzygiella 9
S. subgenus Syzygiella 9
S. autumnalis 9
[S. manca] 69
S. rubricaulis 9
Targionia 27
T. subgenus Targionia 27
T. hypophylla 27
subsp. hypophylla 27
[subsp. linealis] 27
T. lorbeeriana 27
TARGIONIACEAE 7, 27
TAXIPHYLLACEAE 8, 62
Taxiphyllum 62
T. densifolium 62
T. wissgrillii 62
Tayloria 47
T. acuminata 47
T. froelichiana 47
T. hornschuchii 47
T. lingulata 47
T. rudolphiana 47
T. serrata 47
T. splachnoides 47
T. tenuis 47
Taylorioideae 47
Telaranea 19
T. azorica 19
T. europaea 19, 67
T. longii = Tricholepidozia tetradactyla 19
T. murphyae = Tricholepidozia tetradactyla 19
T. nematodes auct. eur. = Telaranea europaea 19
[T. nematodes] 67
T. sejuncta auct. eur. = Telaranea europaea 19
[T. sejuncta] 67
Tetralophozia 10
T. filiformis 10
T. setiformis 10
TETRAPHIDACEAE 7, 29
TETRAPHIDALES 7, 29
Tetraphidopsida 7, 29
Tetraphis 29
T. pellucida 29
Tetraplodon 48
T. angustatus 48
T. blyttii 48
T. mnioides 48
T. pallidus 48
T. paradoxus 48
T. urceolatus 48
Tetrastichium 55
T. fontanum 55
T. virens 55
Tetrodontium 29
T. brownianum 29
T. ovatum 29
T. repandum 29
Thamnobryum 65
T. alopecurum 65
T. angustifolium 65
T. cataractarum 65
T. fernandesii 65
T. maderense 65
T. neckeroides 65
T. rudolphianum 65
T. subserratum 65
THUIDIACEAE 8, 59
Thuidiopsis 59
T. sparsa 4, 59
Thuidium 59
T. assimile 59
T. delicatulum 59
T. recognitum 59
T. tamariscinum 59
Timmia 30
T. section Norvegica 30
T. section Timmia 30
T. section Timmiaurea 30
T. austriaca 30
T. bavarica 30
T. comata 30
T. megapolitana 30
T. norvegica 30
T. sibirica 30
TIMMIACEAE 7, 30
TIMMIALES 7, 30
Timmiella 32
T. anomala 32
T. barbuloides 32
T. flexiseta 32
TIMMIELLACEAE 7, 32
Timmiidae 7
Tomentypnum 58
T. nitens 58
Tortella 42
T. alpicola 42
[T. arctica] 42, 68
T. bambergeri = Tortella tortuosa 43
T. bambergeri auct. p.p. = Tortella fasciculata 43
T. bambergeri auct. p.p. = Tortella pseudofragilis 42
T. x cuspidatissima 42, 68
T. densa 42
T. fasciculata 42
T. flavovirens 42
var. flavovirens 42
JOURNAL OF BRYOLOGY
var. glareicola 42
var. papillosissima 42
T. fragilis 42
var. moravica = Tortella pseudofragilis 42
T. humilis 42
T. inclinata 42
var. densa = Tortella densa 42
T. inflexa 42
T. limbata 42
T. limosella = Tortella flavovirens 42
[T. malacophylla] 70
T. mediterranea 42
T. nitida 42
T. pseudofragilis 42
T. rigens 42
T. spitsbergensis 42
T. squarrosa 42
T. tortuosa 43
subsp. fasciculata = Tortella fasciculata 42
var. fragilifolia = Tortella tortuosa 43
Tortula 39
T. acaulon 39, 43
var. acaulon 39
[var. galilaea] 70
var. papillosa 39
var. pilifera 39
var. retortifolia 39
var. schreberiana 39
T. amplexa 39
T. ampliretis 39
T. atrovirens 39
T. baetica = Tortula israeelis 40
T. bogosica 4, 39
T. bolanderi 4, 39
T. brevissima 39
T. canescens 39
T. caucasica 39
T. cernua 40
T. cuneifolia 40
T. freibergii 40
[T. grandiretis] 70
T. guepinii 40
T. hoppeana 40
T. inermis 40
T. israelis 40
[T. kneuckeri] 70
T. lanceola = Tortula lindbergii 40
T. laureri 40
T. leucostoma 40
T. lindbergii 40
T. lingulata 40
T. marginata 40
T. modica = Tortula caucasica 40
T. mucronifolia 40
T. muralis 40
subsp. muralis 40
subsp. obtusifolia 40
var. aestiva 40
var. muralis 40
T. obtusifolia = Tortula muralis subsp. obtusifolia 40
T. pallida 40
[T. plintobia] 70
T. protobryoides 40
T. randii 40
T. revolvens 40
T. rhodonia = Henediella heimii 38
T. schimperi 40
T. solmsii 40
T. subulata 40
var. angustata = Tortula schimperi 40
var. graefii = Tortula subulata 40
var. subinermis = Tortula subulata 40
T. systylia 40
T. truncata 40
T. ucrainica 40
T. vahliana 40
T. viridifolia 40
T. vlassovii = Microbryum vlassovii 38
T. wilsonii 40
Trachycystis 53
T. ussuriensis 53
Trematodon 36
T. ambiguus 36
T. brevicollis 36
T. laetevirens 36
T. longicollis 36
T. perssoniorum 36
Trichocolea 21
T. tomentella 21
TRICHOCOLEACEAE 6, 21
Trichodon 36
T. cylindricus 36
Tricholepidozia 19
T. lindenbergii 4, 19
[var. complanata] 19
var. lindenbergii 4, 19
[var. mellea] 19
[var. papillata] 19
T. tetradactyla 4, 19
Trichostomoideae 40
Trichostomopsis = Didymodon 37
T. australasiae = Didymodon australiasiae 37
T. umbrosa = Didymodon umbrosus 38
Trichostomum 43
T. arcticum = Tortella spitsbergensis 42
T. brachydontium 43
T. crispulum 43
T. hibernicum = Chionoloma hibernicum 40
T. pallidisetum = Pottiopsis caespitosa 42
T. recurvifolium = Chionoloma recurvifolium 41
T. tenuirostre = Chionoloma tenuirostre 41
T. triumphans = Pottiopsis caespitosa 42
Trilophozia 14
T. quinquedentata 14
[var. assymetrica] 14
var. quinquedentata 14
Triquetrella 40
T. arapilensis 40
Tritomaria 14
T. exsecta 14
subsp. exsecta 14
[subsp. novaezelandia] 14
T. exsectiformis 14
subsp. arctica 14
[subsp. camerunensis] 14
subsp. exsectiformis 14
T. polita = Saccobasis polita 14
subsp. polymorpha = Saccobasis polymorpha 14
T. quinquedentata = Trilophozia quinquedentata 14
subsp. turgida = Trilophozia quinquedentata 14
var. dentata = Trilophozia dentata 14
var. grandiretis = Trilophozia quinquedentata 14
f. gracilis = Trilophozia quinquedentata 14
T. scitula 14
115
116
N. G. HODGETTS ET AL.
Trochobryum = Seligeria 43
T. carniolicum = Seligeria carniolica 43
Tylimanthus = Acrobolbus 16
[T. anisodontus = Acrobolbus anisodontus] 66
T. azoricus = Acrobolbus azoricus 16
T. madeirensis = Acrobolbus madeirensis 16
Ulota 54
U. bruchii 54
U. calvescens 54
U. coarctata 54
U. crispa 54
var. crispula = Ulota crispula 54
var. intermedia = Ulota intermedia 54
U. crispula 54
U. curvifolia 54
U. drummondii 54
U. hutchinsiae 54
U. intermedia 54
U. macrospora 54
U. phyllantha = Phenogemma phyllantha 54
U. rehmannii 54
Vesicularia 63
V. reimersiana 63
Vinealobryum = Didymodon 37
V. brachyphyllum = Didymodon brachyphyllus 37
V. cordatum = Didymodon cordatus 37
V. eckeliae = Didymodon eckeliae 37
V. insulanum = Didymodon insulanus 37
V. luridum = Didymodon luridus 37
V. nicholsonii = Didymodon nicholsonii 38
V. sinuosum = Didymodon sinuosus 38
V. vineale = Didymodon vinealis 38
Voitia 48
V. hyperborea 48
V. nivalis 48
Warnstorfia 58
W. exannulata = Sarmentypnum exannulatum 58
W. fluitans 58
W. procera = Sarmentypnum procerum 58
W. pseudostraminea 58
W. sarmentosa = Sarmentypnum sarmentosum 58
W. trichophylla = Sarmentypnum trichophyllum 58
W. tundrae = Sarmentypnum tundrae 58
Weissia 43
W. angustifolia 43
W. brachycarpa 43
[W. breutelii] 70
W. condensa 43
var. armata 43
var. condensa 43
W. controversa 43
var. controversa 43
var. crispata 43
var. densifolia 43
[W. leptocarpa] 70
W. levieri 43
W. longifolia 43
var. angustifolia = Weissia angustifolia 43
W. x mittenii 43
W. multicapsularis = Tortula acaulon 43
W. multicapsularis auct. = Weissia wilsonii 43
[W. ovatifolia] 70
W. perssonii 43
W. rostellata 43
W. rutilans 43
[W. sinaloensis] 70
W. squarrosa 43
W. sterilis 43
W. triumphans = Pottiopsis caespitosa 42
W. tyrrhena = Pottiopsis caespitosa 42
W. wilsonii 43
W. wimmeriana 43
Zoopsidoideae 19
Zygodon 54
Z. bistratus = Zygodon catarinoi 54
Z. catarinoi 54
Z. conoideus 54
var. conoideus 54
var. lingulatus 54
Z. dentatus 54
Z. forsteri = Codonoblepharon forsteri 53
Z. gracilis 54
Z. rupestris 55
Z. sibiricus 55
Z. stirtonii 55
Z. viridissimus 55
Addenda
1. Herbertus p. 19:
Sun and He (2019) recently proposed synonymising Herbertus borealis, Herbertus norenus and Herbertus stramineus with Herbertus aduncus (in which they include
Herbertus
hutchinsiae). While undoubtedly closely
related, the small genetic differences between them
result in very clear morphological differences, and in the
field the species (which sometimes grow in close proximity) are easily distinguished by morphology, ecology
and distribution. We therefore reject the synonymies.
2. Additional species:
Physcomitrium (Callaghan et al. 2020) p. 31
23.6i Physcomitrium × stevensonii (stevensoni)
D.A.Callaghan
Schistidium p. 46 (Guerra et al. 2020)
122.29i Schistidium memnonium J.Guerra
Meesia (Hedenäs 2020) p. 48
138.4i Meesia minor Brid.
138.4ii Meesia minutissima Hedenäs