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A miniature world
in decline
European Red List of Mosses,
Liverworts and Hornworts
Nick Hodgetts, Marta Cálix, Eve Engleeld, Nicholas Fettes, Mariana García Criado, Lea Patin, Ana Nieto, Ariel Bergamini, Irene
Bisang, Elvira Baisheva, Patrizia Campisi, Annalena Cogoni, Tomas Hallingbäck, Nadya Konstantinova, Neil Lockhart, Marko
Sabovljevic, Norbert Schnyder, Christian Schröck, Cecilia Sérgio, Manuela Sim Sim, Jan Vrba, Catarina C. Ferreira, Olga Afonina,
Tom Blockeel, Hans Blom, Steffen Caspari, Rosalina Gabriel, César Garcia, Ricardo Garilleti, Juana González Mancebo, Irina
Goldberg, Lars Hedenäs, David Holyoak, Vincent Hugonnot, Sanna Huttunen, Mikhail Ignatov, Elena Ignatova, Marta Infante, Riikka
Juutinen, Thomas Kiebacher, Heribert Köckinger, Jan Kučera, Niklas Lönnell, Michael Lüth, Anabela Martins, Oleg Maslovsky,
Beáta Papp, Ron Porley, Gordon Rothero, Lars Söderström, Sorin Ştefǎnuţ, Kimmo Syrjänen, Alain Untereiner, Jiri Váňa Ɨ, Alain
Vanderpoorten, Kai Vellak, Michele Alef, Jeff Bates, Neil Bell, Monserrat Brugués, Nils Cronberg, Jo Denyer, Jeff Duckett, H.J.
During, Johannes Enroth, Vladimir Fedosov, Kjell-Ivar Flatberg, Anna Ganeva, Piotr Gorski, Urban Gunnarsson, Kristian Hassel,
Helena Hespanhol, Mark Hill, Rory Hodd, Kristofer Hylander, Nele Ingerpuu, Sanna Laaka-Lindberg, Francisco Lara, 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 Seneca, Adam Stebel, Risto Virtanen, Henrik Weibull, Jo Wilbraham and Jan Żarnowiec
About IUCN
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A miniature world
in decline
European Red List of Mosses,
Liverworts and Hornworts
Nick Hodgetts, Marta Cálix, Eve Engleeld, Nicholas Fettes, Mariana García Criado, Lea Patin, Ana Nieto, Ariel Bergamini, Irene
Bisang, Elvira Baisheva, Patrizia Campisi, Annalena Cogoni, Tomas Hallingbäck, Nadya Konstantinova, Neil Lockhart, Marko
Sabovljevic, Norbert Schnyder, Christian Schröck, Cecilia Sérgio, Manuela Sim Sim, Jan Vrba, Catarina C. Ferreira, Olga Afonina,
Tom Blockeel, Hans Blom, Steffen Caspari, Rosalina Gabriel, César Garcia, Ricardo Garilleti, Juana González Mancebo, Irina
Goldberg, Lars Hedenäs, David Holyoak, Vincent Hugonnot, Sanna Huttunen, Mikhail Ignatov, Elena Ignatova, Marta Infante, Riikka
Juutinen, Thomas Kiebacher, Heribert Köckinger, Jan Kučera, Niklas Lönnell, Michael Lüth, Anabela Martins, Oleg Maslovsky,
Beáta Papp, Ron Porley, Gordon Rothero, Lars Söderström, Sorin Ştefǎnuţ, Kimmo Syrjänen, Alain Untereiner, Jiri Váňa Ɨ, Alain
Vanderpoorten, Kai Vellak, Michele Alef, Jeff Bates, Neil Bell, Monserrat Brugués, Nils Cronberg, Jo Denyer, Jeff Duckett, H.J.
During, Johannes Enroth, Vladimir Fedosov, Kjell-Ivar Flatberg, Anna Ganeva, Piotr Gorski, Urban Gunnarsson, Kristian Hassel,
Helena Hespanhol, Mark Hill, Rory Hodd, Kristofer Hylander, Nele Ingerpuu, Sanna Laaka-Lindberg, Francisco Lara, 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 Seneca, Adam Stebel, Risto Virtanen, Henrik Weibull, Jo Wilbraham and Jan Żarnowiec
e designation of geographical entities in this book, and the presentation of the material, do not imply the expression of any opinion
whatsoever on the part of the European Commission or IUCN concerning the legal status of any country, territory, or area, or of its authorities,
or concerning the delimitation of its frontiers or boundaries.
e views expressed in this publication do not necessarily reect those of the European Commission or IUCN.
is publication has been prepared by IUCN with support from the IUCN Species Survival Commission and other experts. It is the product of
a LIFE project funded by the European Commission (LIFE14 PRE BE 001).
Project Title: Establishing a European Red List of Bryophytes, Pteridophytes, Saproxylic Beetles, Terrestrial Molluscs and Vasular Plants (LIFE
European Red Lists; LIFE14 PRE BE 001)
Project duration: May 2015 to September 2019
Project total costs: 1,166,667 EUR
Contribution of the LIFE Programme: 700,000 EUR
Co-nancers of the project: National Parks and Wildlife Service, Republic of Ireland; Ministry of Economic Aairs, Department of Nature
& Biodiversity (Ministerie van Economische Zaken, Directie Natuur & Biodiversiteit), the Netherlands; Council of Europe; Oce fédéral de
l‘Environnement, Switzerland; Swedish Environmental Protection Agency (Naturvardsverket), Sweden; British Entomological Society, United
Kingdom; Ministry of Sustainable Development and Infrastructure, Government of the Grand-Duché of Luxembourg.
e LIFE Programme (https://ec.europa.eu/easme/en/life) is the EU’s nancial instrument supporting environmental, nature conservation and
climate action projects throughout the EU. e general objective of LIFE is to contribute to the implementation, updating and development of
EU environmental, nature conservation and climate policy and legislation by conancing projects with European added value.
Published by: IUCN, Brussels, Belgium
© 2019 IUCN, International Union for Conservation of Nature and Natural Resources. All rights reserved. Licenced to the European Union
under conditions.
Reproduction of this publication for educational or other non-commercial purposes is authorised without prior written permission from
the copyright holder provided the source is fully acknowledged. Reproduction of this publication for resale or other commercial purposes is
prohibited without prior written permission of the copyright holder.
Citation: Hodgetts, N., Cálix, M., Engleeld, E., Fettes, N., García Criado, M., Patin, L., Nieto, A., Bergamini, A.,
Bisang, I., Baisheva, E., Campisi, P., Cogoni, A., Hallingbäck, T.,Konstantinova, N., Lockhart, N., Sabovljevic,
M., Schnyder, N., Schröck, C., Sérgio, C., Sim Sim, M., Vrba, J., Ferreira, C.C., Afonina, O., Blockeel, T.,
Blom, H., Caspari, S., Gabriel, R., Garcia, C., Garilleti, R., González Mancebo, J., Goldberg, I., Hedenäs, L.,
Holyoak, D., Hugonnot, V., Huttunen, S., Ignatov, M., Ignatova, E., Infante, M., Juutinen, R., Kiebacher, T.,
Köckinger, H., Kučera, J., Lönnell, N., Lüth, M., Martins, A., Maslovsky, O., Papp, B., Porley, R., Rothero,
G., Söderström, L., Ştefǎnuţ, S., Syrjänen, K., Untereiner, A., Váňa, J. Ɨ, Vanderpoorten, A., Vellak, K., Ale,
M., Bates, J., Bell, N., Brugués, M., Cronberg, N., Denyer, J., Duckett, J., During, H.J., Enroth, J., Fedosov,
V., Flatberg, K.-I., Ganeva, A., Gorski, P., Gunnarsson, U., Hassel, K., Hespanhol, H., Hill, M., Hodd, R.,
Hylander, K., Ingerpuu, N., Laaka-Lindberg, S., Lara, F., Mazimpaka, V., Mežaka, A., Müller, F., Orgaz, J.D.,
Patiño, J., Pilkington, S., Puche, F., Ros, R.M., Rumsey, F., Segarra-Moragues, J.G., Seneca, A., Stebel, A.,
Virtanen, R., Weibull, H., Wilbraham, J. and Żarnowiec, J. (2019). A miniature world in decline: European Red
List of Mosses, Liverworts and Hornworts. Brussels, Belgium: IUCN.
ISBN: 978-2-8317-1993-1 (PDF)
978-2-8317-1994-8 (print)
DOI: https://doi.org/10.2305/IUCN.CH.2019.ERL.2.en
Design and layout: Imre Sebestyén jr. / UNITgraphics.com
Printed by: Media Process S.A., Brussels
Available from: IUCN (International Union for Conservation of Nature)
64 Boulevard Louis Schmidt, 1040 Brussels, Belgium
brussels@iucn.org
www.iucn.org/resources/publications
Picture credits on cover page: Cruet Collar-moss Splachnum ampullaceum (Near reatened moss) © Christian Schröck
All photographs used in this publication remain the property of the original copyright holder (see individual captions for details). Photographs
should not be reproduced or used in other contexts without written permission from the copyright holder.
Printed in Belgium. e text of this book is printed on 135 gsm silk paper (300 gsm silk for the cover). is book is printed on recycled paper
made from wood bre from well-managed forests certied in accordance with the rules of the Forest Stewardship Council (FSC).
Ɨ Deceased
iii
Contents
Foreword ........................................................................................................................................................... iv
Preface ................................................................................................................................................................ v
Acknowledgements ............................................................................................................................................ vi
Executive summary ........................................................................................................................................... vii
Recommendations ...........................................................................................................................................viii
1. Background ....................................................................................................................................................1
1.1 e European context .....................................................................................................................................................1
1.2 European mosses, liverworts and hornworts ...................................................................................................................1
1.3 Assessment of species extinction risk ...............................................................................................................................7
1.4 Objectives of the assessment ........................................................................................................................................... 7
2. Assessment methodology ...............................................................................................................................9
2.1 Geographic scope ............................................................................................................................................................ 9
2.2 Taxonomic scope ............................................................................................................................................................. 9
2.3 Assessment protocol ........................................................................................................................................................9
2.4 Species mapping ............................................................................................................................................................ 10
3. Results .........................................................................................................................................................11
3.1 reat status .................................................................................................................................................................. 11
3.2 Status by taxonomic group ...........................................................................................................................................11
3.3 Spatial distribution of species .........................................................................................................................................14
3.4 Major threats to moss, liverwort and hornwort species in Europe .................................................................................. 18
3.5 Population trends .........................................................................................................................................................22
3.6 Gaps in knowledge ........................................................................................................................................................ 23
4. Conservation actions .................................................................................................................................... 26
4.1 Conservation of moss, liverwort and hornwort species in Europe .................................................................................. 26
4.2 Red List versus priority for conservation action .............................................................................................................31
5. Recommendations .......................................................................................................................................32
5.1 Recommended actions ...................................................................................................................................................32
5.2 Application of project outputs ....................................................................................................................................... 33
5.3 Future work .................................................................................................................................................................. 33
References ......................................................................................................................................................... 35
Appendix 1. List of lead assessors by geographical region ................................................................................. 39
Appendix 2. Example of species summary and distribution map ...................................................................... 39
Appendix 3. Red List status of European mosses, liverworts and hornworts ..................................................... 45
Appendix 4. Listing of bryophyte species under Annex II and Annex V of the Habitats Directive,
Appendix I of the Bern Convention, and the Red List status on the current European Red List ....................... 85
IUCN Red List of reatened Species™ – European Regional Assessment Reports ........................................... 87
iv
Foreword
Europe has a rich natural heritage, with habitats ranging
from dry Mediterranean maquis in the south to the Arctic
tundra in the far north. Much of Europe’s landscape has
been shaped by centuries of diverse farming and forestry
traditions. As a result, a large number of agricultural and
semi-natural landscapes have emerged and signicantly
inuenced the continent’s biodiversity.
Biodiversity loss as a result of human impacts is one of the
major challenges that the world currently faces, and this
has considerably aected valuable ecosystem services. In
order to halt the loss of biodiversity, the EU Biodiversity
Strategy aims to protect, value and restore biodiversity
and the services it provides – Europe’s natural capital.
is is important not only to protect nature for its own
sake, but also for its essential contribution to human well-
being and economic prosperity, and to avert catastrophic
changes caused by biodiversity loss. After all, everyone
will understand that we cannot act sustainably if we keep
destroying nature.
In recent years, awareness has risen surrounding the
crucial role of plants in providing ecosystem services
and on their decline – they are one of the essential
foundations of healthy ecosystems that we depend on.
However, signicant gaps in knowledge still remain. In
this context, A miniature world in decline: European Red
List of Mosses, Liverworts and Hornworts provides the
rst-ever comprehensive assessment of the extinction risk
of all native bryophyte species to Europe. With 1,817
species assessed, this assessment highlights that 22.5%
of bryophyte species are threatened with extinction
in Europe. Main threats to these species include land
modications such as those caused by the construction
of dams and through increased frequency of res, the
impacts of climate change as well as agricultural and
forestry practices, such as the conversion of natural
forest to plantation woodland. While 88.2% of species
are recorded in at least one protected area (whether it
is a national park, Natura 2000 site or nature reserve),
eight species are already lost from Europe, and so it is
clear that we continue to have a responsibility to conserve
these unique species to prevent further extinctions in our
region.
By comparison to European bryophytes, 2% of medicinal
plants, 8% of aquatic plants, 16% of crop-wild relatives,
20% of ferns and lycopods, 42% of trees and 57% of
“policy plants” (listed under European or international
policy instruments) are threatened, ranking bryophytes
third as the most threatened group of plant species
assessed so far.
Immediate action must be taken in order to improve
the status of European bryophytes. A multidisciplinary
approach needs to be established; while species and
protected area management are key, the proper
implementation of the existing European legislation
will be crucial in providing protection to the species,
including the EU Birds and Habitats Directive.
I hope that this new IUCN European Red List will
help place plants higher on the conservation agenda
as well as inform the wider debate and contribute to
the discussion on priorities within the conservation
community. A network of bryophyte experts is already
in place, therefore more investment in scientic research,
and increasing awareness and communications will help
towards the delivery of real results and positive impacts
for these species.
Humberto Delgado Rosa
Director for Natural Capital
DG Environment, European Commission
v
Preface
is publication has been prepared by IUCN
(International Union for Conservation of Nature) as
a deliverable of the LIFE European Red Lists project
(LIFE14 PRE BE 001). A miniature world in decline: e
European Red List of Mosses, Liverworts and Hornworts is,
therefore, a part of a series of publications released since
2015, when the project began, that also include:
• European Red List of Lycopods and Ferns, 2017
• European Red List of Saproxylic Beetles, 2018
• European Red list of Terrestrial Molluscs: slugs, snails,
and semi-slugs, 2019
• European Red list of Trees, 2019
• European Red list of Selected Endemic Shrubs, 2019
Based on other European Red List assessments, 59%
of freshwater molluscs, 40% of freshwater shes, 28%
of grasshoppers, crickets and bush-crickets, 23% of
amphibians, 20% of reptiles, 20% of ferns and lycopods,
17% of mammals, 16% of dragonies, 13% of birds,
9% of butteries and bees, 8% of aquatic plants and 2%
of medicinal plants are threatened at the European level
(Allen et al., 2014; IUCN, 2015; Hochkirch et al., 2016;
García Criado et al., 2017). Additional European Red
Lists assessing a selection of species showed that 22% of
terrestrial molluscs, 16% of crop wild relatives and 18%
of saproxylic beetles are also threatened (Cuttelod et al.,
2011; Bilz et al., 2011; Cálix et al., 2018). e ndings of
this work suggest that 23% of bryophytes are threatened
species in Europe, representing the fth most threatened
group of plants assessed so far.
Lindenberg’s featherwort Adelanthus lindenbergianus (Endangered liverwort) © Rory Hodd
vi
Acknowledgements
All of IUCN’s Red Listing processes rely on the willingness
of scientists to contribute and pool their collective
knowledge to make the most reliable estimates of the
status of a species. Without their enthusiastic commitment
to species conservation, this kind of regional overview
would not be possible. Bryophytes are no exception,
and the knowledge mobilized through the Europe-wide
network of members of the European Committee for the
Conservation of Bryophytes (ECCB) has been pivotal to
the completion of this Red List. We are therefore indebted
for their support and contributions.
anks go to the Red List Unit, in particular Caroline
Pollock, David Allen, Jemma Window, Kate Harding and
Anna Puttick for their support in the coordination of the
European Red List of Mosses, Liverworts, and Hornworts.
Anna Rosenberg and Corinna Karlsen provided substantial
assistance with nancial management of the project.
A word of thanks goes to Marc Hall for his support on
communication-related matters. We are also grateful to
the many photographers, who provided photos for the
species assessments and the photos in this publication.
We also thank Craig Hilton-Taylor and Malin Rivers for
peer-reviewing this publication.
e European Red List of Mosses, Liverworts and
Hornworts, and consequently this publication, are
required as part of a LIFE project funded by the European
Commission and other donors (LIFE14 PRE BE 001 –
LIFE European Red Lists). In particular, we would like
to thank Frank Vassen (European Commission) and the
LIFE monitors, particularly Kristijan Civic, for their
support throughout the project, allowing for a smooth
implementation.
vii
Executive summary
Aim
is Red List is a summary of the conservation status of
the European species of mosses, liverworts and hornworts,
collectively known as bryophytes, evaluated according
to IUCN’s Guidelines for Application of IUCN Red
List Criteria at Regional Level. It provides the rst
comprehensive, region-wide assessment of bryophytes
and it identies those species that are threatened with
extinction at a European level, so that appropriate policy
measures and conservation actions, based on the best
available evidence, can be taken to improve their status.
Scope
All bryophytes native to or naturalised in Europe (a total
of 1,817 species), have been included in this Red List. In
Europe, 1,796 species were assessed, with the remaining
21 species considered Not Applicable (NA). For the EU
28, 1,728 species were assessed, with a remaining 20
species considered NA and 69 species considered Not
Evaluated (NE). e geographical scope is continent-
wide, extending from Iceland in the west to the Urals in
the east, and from Franz Josef Land in the north to the
Canary Islands in the south. e Caucasus region is not
included. Red List assessments were made at two regional
levels: for geographical Europe and for the 28 Member
States of the European Union.
Results
Overall, 22.5% of European bryophyte species assessed in
this study are considered threatened in Europe, with two
species classied as Extinct and six assessed as Regionally
Extinct (RE). A further 9.6% (173 species) are considered
Near reatened and 63.5% (1,140 species) are assessed
as Least Concern. For 93 species (5.3%), there was
insucient information available to be able to evaluate
their risk of extinction and thus they were classied as Data
Decient (DD). e main threats identied were natural
system modications (i.e., dam construction, increases
in re frequency/intensity, and water management/use),
climate change (mainly increasing frequency of droughts
and temperature extremes), agriculture (including
pollution from agricultural euents) and aquaculture.
Exormotheca welwitschii (Endangered liverwort) © Michael Lüth
viii
Recommendations
Policy measures
• Use the European Red List as the scientic basis to
inform regional/national lists of rare and threatened
species and to identify priorities for conservation
action in addition to the requirements of the Habitats
Directive, thereby highlighting the conservation
status of bryophytes at the regional/local level.
• Use the European Red List to support the integration
of conservation policy with the Common Agricultural
Policy (CAP) and other national and international
policies. For example, CAP Strategic Plans should
include biodiversity recovery commitments that could
anticipate, among others, the creation of Important
Bryophyte Areas. An increased involvement of
national environmental agencies in the preparation
of these strategic plans, and more broadly in ongoing
discussions on the Future CAP Green Architecture,
would likely also ensure the design of conservation
measures better tailored to conserve bryophytes in
agricultural landscapes.
• Update the European Red List every decade to ensure
that the data remains current and relevant.
• Develop Key Biodiversity Areas for bryophytes in
Europe with a view to ensuring adequate site-based
protection for bryophytes.
Research and monitoring
• Use the European Red List as a basis for future tar-
geted eldwork on possibly extinct and understudied
species.
• Establish a monitoring programme for targeted spe-
cies (for example, threatened species and/or arable
bryophytes).
• Use the European Red List to obtain funding for re-
search into the biology and ecology of key targeted
species.
Action on the ground
• Use the European Red List as evidence to support
multi-scale conservation initiatives, including
designation of protected areas, reform of agricultural
practices and land management, habitat restoration
and rewilding, and pollution reduction measures.
• Use the European Red List as a tool to target species
that would benet the most from the widespread
implementation of the solutions oered by the
1991 Nitrates Directive (Council Directive 91/676/
EEC), including the application of correct amounts
of nutrients for each crop, only in periods of crop
growth under suitable climatic conditions and never
during periods of heavy rainfall or on frozen ground,
and the creation of buer zones to protect waters
from run-o from the application of fertilizers.
Ex situ conservation
• Undertake ex situ conservation of species of
conservation concern in botanic gardens and spore
and gene banks, with a view to reintroduction where
appropriate.
1
1. Background
1.1 The European context
Europe is the world’s second smallest continent in terms
of area after Australia, covering approximately 10.4
million km², or 2% of the Earth’s surface. In terms of
human population, Europe is the third largest continent
(after Asia and Africa) with a population of around 546
million (UN DESA, 2018) – about 13% of the world’s
population. erefore, Europe is one of the smallest and
one of the most densely populated continents in the
world.
e European Union (EU), consisting of 28 Member
States (EU 28), is Europe’s largest political and economic
entity. e ecological footprint of the EU 28 has been
estimated to exceed the region’s biological capacity (the
total area of cropland, pasture, forest, and shing grounds
available to produce food, bre and timber, and absorb
waste) by 2.6 times (EEA, 2015).
Europe has a great diversity of landscapes and habitats
and a wealth of ora and fauna. For example, the
Mediterranean Basin, which is especially rich in plant and
animal species, many of them endemic to that region,
has been recognised as a global biodiversity hotspot
(Mittermeier et al., 2004; Cuttelod et al., 2008).
e European continent has a highly fragmented
landscape, and up to 80% of land in Europe is currently
used for settlement, industry, production systems
(including agriculture and forestry) and infrastructure
(EEA, 2006; Pedroli & Meiner, 2017). Consequently,
European species are to a large extent dependent upon
habitats created and maintained by human activity, and
many are aected by overexploitation, pollution and the
impacts of invasive alien species. Additionally, climate
change is becoming an increasingly serious threat. Europe
is a diverse region and the relative importance of dierent
threats varies widely across its biogeographic regions and
countries.
Although considerable eorts have been made to protect
and conserve European habitats and species (see Sections
4.1 and 4.2), and the Natura 2000 network of protected
areas covers more than 18% of the EU terrestrial
territory, biodiversity decline and the associated loss
of vital ecosystem services (such as water purication,
pollination, ood protection and carbon sequestration)
continues to be a major concern in the region.
1.2 European mosses, liverworts and
hornworts
Bryophytes are a large, diverse group of plants. According
to Villareal et al. (2010), there are between 18,000 and
23,000 described species worldwide, comprising about
11,000-13,000 mosses, 7,000-9,000 liverworts and 200-
250 hornworts, making them second only to owering
plants in terms of species richness. is could, however,
be an underestimate, with molecular studies revealing
‘new’ species all the time. Each of the three groups of
bryophytes has been traditionally considered to be a
separate phylum (or division): Bryophyta (mosses),
Marchantiophyta (liverworts) and Anthocerotophyta
(hornworts) (Frey & Stech, 2009), although the latest
evidence, with increasing support, suggests that mosses
and liverworts form a clade, termed “Setaphyta” (Puttick
et al., 2018). Nevertheless a number of biological and
ecological characters are common to the three groups:
ey are small (rarely larger than a few centimeters),
unable to produce lignin (they cannot become woody),
have their life cycle dominated by the gametophyte
(rather than the sporophyte) generation – see Box 1 - and
are able to dry out completely in dry periods, quickly
resuming their metabolism when rewetted. ey fulll a
range of important ecological functions, particularly in
water retention, soil-building and in their relationships
with other organisms. For example, bog-moss (Sphagnum
spp.) is one of the most important plants, and certainly
the most important peat producer in the world, locking
away an enormous amount of carbon and holding vast
quantities of water: bogs are essentially huge sponges.
Bryophytes, particularly epiphytes, are also great
indicators of air pollution. Bryophytes show a vast range
of specic sensitivity and visible symptoms to pollutants
greatly exceeding that of higher plants (Govindapyari et
al., 2010).
Mosses
e most species-rich of the three main groups of
bryophytes, mosses, encompasses a wide range of forms.
‘Typical’ mosses (class Bryopsida) are mostly small,
2
Box 1 - The life history of bryophytes
What distinguishes bryophytes collectively from all other land plants is that their life cycle is dominated by the
gametophyte generation; that is, by the haploid or sexual phase, as opposed to the diploid, spore-producing
phase. In contrast, all owering plants, conifers and ferns are dominated by the sporophyte generation, with the
gametophyte much reduced, often to just a few cells. In other words, the main plant that one sees, the leafy green
part, that is mainly photosynthetic, is the gametophyte in bryophytes, whereas it is the sporophyte in all other
plants. e bryophyte sporophyte is usually reduced to a spore-producing, stalked capsule that remains attached
to the gametophyte, and is entirely dependent on it for sustenance.
Figure 1. e life cycle of a bryophyte © MADBRYO initiative
e spore is the rst stage in the haploid gametophyte generation, with a single set of unpaired chromosomes.
Spores germinate into a green protonema, from which the mature gametophyte grows, either producing a
structure with a stem and leaves (as in mosses and leafy liverworts) or a structure with no dierentiation, usually
a at plate of tissue called a thallus (as in thallose liverworts and hornworts). e gametophyte produces (either
on the same plant [bisexual, Figure1] or on separate plants [unisexual, Fig. 1]) male and female sex organs
(antheridia and archegonia, respectively). e antheridia produce sperm, which swim in a lm of water to
the archegonia. In each archegonium only a single egg cell is found which may be fertilized by a sperm cell. If
fertilization has been successful, i.e. if male and female elements of the gametophyte have fused, a new sporophyte
starts to develop. e sporophyte relies largely on the gametophyte for its nutrition. At maturity it eventually
produces spores by the process of meiosis, the kind of cell division that halves the number of chromosomes. In
other words, the brief reign of the sporophyte is over, and a new gametophyte generation is ready to develop.
While there is a lot of variation in the detail of how bryophytes conduct their life cycle, they all conform to
this basic pattern. In addition, many of them produce specialised asexual reproductive organs, such as gemmae,
which circumvent the sporophyte generation entirely, simply replicating the gametophyte parent. In addition,
all bryophytes are to some extent totipotent: they can regenerate from fragments, or even single cells, making
them great survivors.
3
rather delicate, often translucent plants that absorb
water and nutrients externally, over their entire surface.
e sporophyte consists of a capsule in which the spores
are produced supported by a stalk (seta). When mature,
the capsule releases its spores through an opening at the
top, which is usually surrounded by a ring of tooth-like
structures (the peristome).
Not all mosses conform to this general template. ere
are a few groups of mosses that are so distinctive they
are put in their own classes within the Bryophyta. Most
obviously, and certainly most importantly, are the bog-
mosses, the genus Sphagnum (class Sphagnopsida) with
61 species in Europe. ey dier from typical mosses
in almost every respect, except for the dominance of
the gametophyte generation. Its unique cell structure
allows Sphagnum to take up water quickly by capillary
action, and release it only very slowly, like a sponge.
Other features of its physiology and morphology make
it possible for Sphagnum to dominate entire landscapes,
as in the extensive boglands of northern Europe, which
store tremendous amounts of peat built by thousands of
years of Sphagnum growth.
A further oddity among the mosses is Andreaea (rock-
moss, class Andreaeopsida), tiny black or very dark
red-brown tufts on acid rocks in the mountains, with
a capsule that splits into four lobes joined at the top,
a bit like a miniature Japanese lantern. en there are
the haircap mosses, the genus Polytrichum and its allies
(class Polytrichopsida). As well as having a distinctive
spore capsule, these plants have an internal conduction
system somewhat analogous to the xylem and phloem
of vascular plants, enabling them to grow much bigger
than ‘ordinary’ mosses, and shoots of Polytrichum are
capable of attaining heights of half a metre or even more.
A) Five-ranked Bog-moss Sphagnum quinquefarium (Least Concern moss) © Christian Schröck, B) snow rock-moss Andreaea nivalis (Near reatened moss) © Michael Lüth,
C) Polytrichum commune (Least Concern moss) © Fred Rumsey, D) Oedipodium grithianum (Near reatened moss) © Vladimiar Fedosov
A
C
B
D
4
Another couple of small classes, the Oedipodiopsida
and the Tetraphidopsida, dier fundamentally from the
Bryopsida in features of the sporophyte, but have few
species.
Liverworts
A less species-rich group than mosses, yet showing a
greater range of forms, liverworts can be subdivided
into leafy (class Jungermanniopsida) and thallose (class
Marchantiopsida), plus a group of rather anomalous plants
that show features of both leafy and thallose liverworts
(class Haplomitriopsida, with only a single species in
Europe).
Leafy liverworts are especially diverse in the form of the
leaves, which range from entire and rounded, through
simply bilobed or trilobed, to deeply divided into
laments or asymmetrically divided so that one lobe is
larger than the other, or even modied into a tiny pocket
or helmet-shaped structure. Furthermore, while leafy
liverworts typically have two rows of main (lateral) leaves
running down opposite sides of the stem, many species
have an additional row of leaves on the under-surface of
the stem (underleaves, or amphigastria): these are usually
smaller than the lateral leaves, and often quite dierent
in shape.
e form of the sporophyte is relatively uniform among
the liverworts, although there is great variation in
the various gametophytic structures that support and
protect it. Unlike mosses, where the spore capsule and
seta mature slowly together, and then spores are released
gradually, the liverwort capsule matures inside a protective
sheath (usually a structure derived from modied leaves
called a perianth), and is raised up on a seta only when
ready to release its spores. e seta therefore grows at a
tremendously fast rate, by sudden elongation of its cells,
and forms a delicate, ephemeral structure which lasts just
long enough for the mature capsule to release all its spores
at once.
Hornworts
e least species-rich group of bryophytes, the hornworts,
supercially resemble thallose liverworts, but are not
closely related. ey probably emerged as a group at about
the same time as the other bryophytes, in the Ordovician
period about 470 million years ago, or even earlier (Morris
et al., 2018), but whether they were even then part of
the same taxonomic group as other bryophytes is still a
matter for research. ey have an unusual combination
of features, some shared with other bryophytes, some
having more in common with vascular plants or algae. For
example, the thallus cells contain just one large chloroplast,
while the sporophyte, which is a long narrow structure
with no dierentiation into seta and capsule, has stomata.
Hornworts reach their greatest diversity in the tropics,
with only a small handful of species occurring in Europe.
Nonetheless, certain species can be locally abundant given
the right conditions.
Distribution, habitats and ecology
Bryophytes occur on all continents and in many dierent
habitats except in the sea. ey are almost ubiquitous,
growing even in very dry semi-deserts, but require some
moisture, at least at some stages of their life cycle. Unlike
vascular plants, most species are poorly equipped to
regulate their water content internally, instead drying out
and rewetting rapidly according to external conditions
A) Mannia triandra (Vulnerable liverwort) © Christian Schröck, B) Phaeoceros carolinianus (Near reatened hornwort) © Michael Lüth
A B
5
(poikilohydric). is means that they are often luxuriant in
moist forest and in high rainfall areas. Bryophytes absorb
water, along with the minimal amounts of nutrients they
require, over their entire surface from the surrounding
environment, rather than taking it up through roots and
a vascular system. However, many bryophytes, and most
of the European bryophytes, have a physiology that allows
them to dry out completely in the absence of moisture,
suspend physiological activity, and then ‘come back
to life’ when wetted again. Dierent species do this to
dierent degrees, but herbarium specimens of the great
hairy screw-moss (Syntrichia ruralis) were recently found
to have retained their vitality after over 20 years dried in a
packet (Stark et al., 2016), and there are unsubstantiated
anecdotes about much longer periods of survival in the
herbarium. Many species which grow directly on rock
in exposed conditions (Grimmia, Didymodon, etc.) dry
out and rehydrate virtually on a daily basis, particularly
in warmer climates; this is their strategy for enduring
drought. is contrasts with most vascular plants, which
could not survive this level of dehydration.
Bryophytes have several ecological attributes that are very
distinctive:
• ey are poikilohydric, i.e., they dry out and rewet
rapidly according to external conditions.
• ey grow in ‘microhabitats’: whether they grow in
woodland, heathland or grassland is less important
than the immediate micro-environment, such as a rock
crevice or a moist patch of soil.
• A large proportion of species are colonists, and therefore
form pioneer communities and assemblages. ere are,
for example, many short-lived ruderal (weedy) species
that colonise bare ground, disappearing as vegetational
succession proceeds.
• Many species are very ecient at dispersal, with
spores and vegetative propagules potentially capable of
travelling worldwide in the air or via vectors such as
migratory birds.
• Due to the latter, levels of endemism are low (in Europe,
ca. 10% compared to ca. 28% vascular plants; Patino
& Vanderpoorten, 2018), but levels of disjunction
are high (for example, a species may occur in western
Europe, the Himalayas and British Columbia).
• ose species that are less ecient at dispersal often
have large spores that can remain viable in the soil for
a long time.
• ey are often excellent ecological indicators (for
example, of nutrient status or pH)
.
Ecosystem services and commercial use
e ecosystem services that bryophyte species provide
might not be conspicuous, but investigation soon shows
us that these small plants, useless as food or building
materials, are actually of vital importance. In particular,
there are three main features of bryophytes that make
them important in the ecosystem:
Grimmia mollis (Vulnerable moss) © Michael Lüth
6
• eir ability to retain water. All bryophytes act to
some extent as sponges, taking up water rapidly,
holding it, and releasing it only slowly. is is most
obvious in bog-moss Sphagnum, which dominates
vast areas of mire in northern Europe. On hillsides
and hilltops, Sphagnum is an important stabilising
inuence in areas with heavy rainfall. A similar eect
is seen in forest ecosystems. A substantial part of the
water-holding capacity of forests is bound up in the
bryophytes, and when it is clear-felled, the resultant
erosion, ooding and destabilisation is at least partly
because the bryophytes have been removed from the
landscape along with the trees.
• ey are ecient colonisers and stabilisers of bare
substrates. When natural erosion occurs, bryophytes
are usually the rst plants to appear on the newly
exposed surfaces. After volcanic eruptions, bryophytes
are the rst to colonise the cooling lava ows. In post-
industrial landscapes, and in urban habitats more
generally, bryophytes often build up thick carpets over
crumbling concrete and tarmac, trapping detritus,
building new soils, providing rooting substrates for
larger plants and ultimately the basis for entire new
ecosystems.
• ey serve as hosts for blue-green algae (cyanobacteria),
which have an important role in nitrogen (N)
xation, and provide a major source of N for boreal
ecosystems (for example, Ackerman, 2013). It is
likely that epiphytic cyanobacteria are a key factor in
determining the abundance of feather mosses across
the boreal biome (Zacckrissson et al., 2019).
Additionally, they provide habitats for other organisms;
seed-beds for vascular plants, shelter and food for
invertebrates, nesting material for birds and small
mammals. Bogs in particular form entire ecosystems
fundamentally dependent on bryophytes.
Sphagnum mosses are used commercially - peat is (or
has been, historically) burned for fuel. However, this
has rarely been done in a sustainable way, and several
countries have established peat-burning power stations,
which have had a devastating eect on peatlands. It has
also been used as a mildly antiseptic dressing for wounds
(it was harvested for this purpose during the First World
War), and as an absorbent material in babies’ nappies.
Bryophytes are harvested commercially (sustainably or
otherwise) for horticultural purposes such as packing
material for bulbs or a water-retentive substrate for
hanging baskets. Sphagnum harvested for horticultural
purposes is also not usually sustainable. A more modern
use is that of mosses, particularly the Habitats Directive
e procession of ‘moss men’ in Béjar, Spain, commemorating the use of moss as camouage in battles during the 12th century by local Christians © Eloy Díaz-Redondo
7
Annex V Leucobryum, for ‘moss walls’ and other ‘green’
architectural purposes. Other commercial uses are largely
historical or minor: Polytrichum commune for brooms,
mosses as insulation for homes, decorative garlands and
even clothing for the famous ‘moss men’ of Béjar in Spain.
1.3 Assessment of species extinction risk
e conservation status of plants, animals and fungi is
one of the most widely used indicators for assessing the
condition of ecosystems and their biodiversity. At the
global scale, the primary source of information on the
extinction risk of plants and animals is e IUCN Red
List of reatened SpeciesTM (www.iucnredlist.org), which
contributes to understanding the conservation status of
assessed species.
e IUCN Red List Categories and Criteria (IUCN,
2012a) are designed to determine the relative risk
of extinction of a taxon, with the main purpose of
cataloguing and highlighting those taxa that are facing a
high risk of extinction. e IUCN Red List Categories
are based on a set of quantitative criteria linked to
population trends, size and structure, threats, and
geographic ranges of species. When conducting regional
or national assessments, the IUCN Red List Regional
Guidelines (IUCN, 2012b) are applied to assign the
IUCN Red List Categories (Figure 2).
As the extinction risk of a species can be assessed at global,
regional or national levels, a species may have a dierent
Red List Category on the global Red List than on the
regional Red List. Logically, an endemic species should
have the same Category at regional and global levels, as it
is not present anywhere else in the world.
1.4 Objectives of the assessment
e European Red List of Mosses, Liverworts and
Hornworts has four main objectives:
• to contribute to regional conservation planning
through provision of a baseline dataset reporting the
conservation status of European bryophyte species;
• to identify those priority geographic areas and habitats
needing to be conserved to prevent extinctions and to
ensure that European bryophytes reach and maintain
a favourable conservation status;
• to identify the major threats and to propose potential
mitigating measures and conservation actions to
address them;
• to strengthen the network of experts focused on
bryophyte conservation in Europe, so that the
assessment information can be kept current and
expertise can be targeted to address the highest
conservation priorities.
Figure 2. e IUCN Red List Categories at the regional scale.
8
e assessment provides three main outputs:
• summary reports on the status of all 1,817 European
bryophyte species;
• a freely available database holding the baseline data on
the status and distribution of European bryophytes;
• a website and data portal (http://ec.europa.eu/
environment/nature/conservation/species/redlist and
www.iucnredlist.org/initiatives/europe) showcasing
these data in the form of species factsheets for all
European bryophytes included in this study, along
with background and other interpretative material.
• is Red List provides the rst comprehensive,
region-wide assessment of bryophytes and builds on
the previous work of the European Committee for the
Conservation of Bryophytes (ECCB). e enormous
amount of new eldwork, data and knowledge
accumulated since then means that it should be much
more robust and authoritative. Eorts will continue
to update the database which will also be made freely
and widely available.
Matted bryum Bryum calophyllum (Endangered moss) © Neil Lockhart
9
2. Assessment methodology
2.1 Geographic scope
e geographic scope is continent-wide, extending from
Iceland in the west to the Urals in the east (including
European parts of the Russian Federation), and from
Franz Josef Land in the north to the Mediterranean in the
south (Figure 3). e Canary Islands, Selvagens, Madeira,
the Azores, Malta and Cyprus are also included. In the
southeast, the Caucasus region and Anatolia are excluded.
Red List assessments were made at two regional levels:
1) for geographical Europe (limits described above); and
2) for the area of the 28 Member States of the European
Union (EU 28) (as of 2018).
2.2 Taxonomic scope
e European Red List of Mosses, Liverworts and
Hornworts has assessed the status of all bryophyte species
considered native to or naturalised in Europe. e original
list of species was based on Hodgetts (2015), which was
in turn based on Hill et al. (2006) for the mosses and
Söderström et al. (2007) for the liverworts and hornworts.
e inclusion of newly described or species which have
undergone taxonomic change (up to the end of 2018)
was undertaken following consultation with the relevant
experts. When there were discrepancies in the identity of
a species, consultation was sought among the dierent
specialists and decisions were made through consensus.
2.3 Assessment protocol
For all the bryophyte species assessments, the following
data were compiled:
• taxonomic classication and notes
• geographic range and list of countries of occurrence
(including a range map)
Figure 3. European assessment boundaries: regional assessments were made for two areas: geographical Europe
and the EU 28.
10
• population information and overall population trends
• habitat preferences and primary ecological
requirements, including pertinent biological
information (for example, generation length,
maximum size and age, etc.)
• species use and trade
• major threats
• research needs
• conservation measures (in place and needed)
• IUCN Red List Category and Criteria and rationale
• key literature references
Some critical terms like ‘mature individual’, ‘generation
length’, and ‘severely fragmented’ had to be interpreted
in a pragmatic way so that they became applicable
to bryophytes. Work over several years (for example,
Hallingbäck et al., 1998) and collaboration with the
IUCN under this project, has culminated in a paper
addressing these issues (Bergamini et al., 2019)
1
.
e task of collecting the initial data was divided
geographically between 11 Assessors (Appendix 1), and
information on each species was based on published and
unpublished data and expert knowledge. e IUCN
Species Information Service (SIS) was used to enter and
store all species data.
A training workshop was held in October 2015 in Paris
(France) in order to train the experts on the IUCN Red
List methodology. After the preliminary information was
collected by the Lead Assessors, ve assessment workshops
were held to review and discuss the assessments and
distribution maps, add new information to the assessments,
and agree on the nal IUCN Red List Category and
Criteria for each species. e workshops took place at the
Faculty of Sciences of the University of Lisbon (Portugal;
December 2016), the Ministry of Environment of the
Czech Republic (Prague; January 2017), the IUCN
European Regional Oce in Brussels (Belgium; February
2017), the National Botanical Gardens of Ireland
(Dublin; April 2017), and ArtDatabanken at the Swedish
University of Agricultural Sciences (Uppsala, June 2017).
In addition, some discussions on the methodology were
held at an external workshop in Ekenäs (Sweden). Overall,
60 experts participated in the assessment workshops.
1 e recommendations from this paper will need to be formally considered
by the IUCN SSC Red List Standards and Petitions Committee, and if
approved, incorporated into a future update of the Red List Guidelines.
For the purposes of this project and for the publication of the assessments
on the IUCN Red List website, the modied approach has been
provisionally approved.
Following the workshops, the information was edited
and any remaining questions were resolved through
communications with the Lead Assessors. An additional
peer-review process was carried out, with all assessments
checked by external Reviewers who had not been previously
involved in the assessment process. Consistency in the
use of IUCN Categories and Criteria was systematically
checked by IUCN sta. e resulting nalised IUCN
Red List assessments are a product of scientic consensus
concerning species status and are supported by relevant
literature and data sources (see example in Appendix 2).
e nal list of species is found in Appendix 3.
2.4 Species mapping
Distribution data were mainly obtained from published
literature, herbarium specimens, internet sources (for
example, GBIF) and several global and regional citizen
science projects. e species experts provided the
distribution data to the Ministry of Environment of the
Czech Republic (MZP) where Jan Vrba compiled the data
in order to produce the nal distribution maps.
Range maps were created using the distribution data
available, which varied in terms of quality; for some
regions, distributional data were available as point locality
data (latitude/longitude) or in grid cell format, and were
therefore spatially precise. Where point or grid data
were available, these were projected in a Geographical
Information System (GIS) (ESRI ArcMap). Polygons
were then drawn manually, clustering occurrence data
where appropriate. In some rare cases where no point data
was available and it was only possible to assign presence
at the country level, the distribution was mapped for the
whole country.
e spatial analyses presented in this publication (see
section 3.3) were done using a geodesic discrete global
grid system, dened on an icosahedron and projected to
the sphere using the inverse Icosahedral Snyder Equal Area
(ISEA) Projection (S39). is corresponds to a hexagonal
grid composed of individual units (cells) that retain their
shape and area (864 km²) throughout the globe. ese
are more suitable for a range of ecological applications
rather than the most commonly used rectangular grids
(S40). e known current distributions (IUCN, 2014) of
extant and possibly extant species were converted to the
hexagonal grid for the purposes of the analysis. Coastal
cells were clipped to the coastline.
11
3. Results
3.1 Threat status
At the European level, 22.5% of bryophyte species are
considered threatened (i.e., assessed as having an elevated
risk of extinction). However, the proportion of threatened
species is uncertain given the number of Data Decient
(DD) species and could lie between 21.4% (if all DD
species are not threatened) and 26.6% (if all DD species
are threatened) for Europe (IUCN, 2011; Table 1). e
mid-point gure provides the best estimation of the
proportion of threatened species (IUCN, 2011). In the
EU 28, 24.3% of species are considered to be threatened,
with the proportion of threatened species lying between
23.1% (if all DD species are not threatened) and 27.9%
(if all DD species are threatened, Table 1). Appendix 3
provides an exhaustive list of all bryophyte species assessed
under the current European Red List and corresponding
conservation status in Europe, EU28 also indicating if the
species is endemic or not to Europe.
In Europe, six species (0.3%) are assessed as Regionally
Extinct, with two endemic species assessed as Extinct
(0.1%). 59 species (3.3%) are Critically Endangered, 143
species (8%) are Endangered, and 180 species (10%) are
Vulnerable (Table 2). A further 173 species (9.6%) are
classied as Near reatened. For 93 species (5.2%) there
were insucient data to evaluate their risk of extinction
and so they were classied as Data Decient (Table 2,
Figure 4). ere were 21 species that were classed as Not
Applicable in Europe (species introduced after AD 1500
or species of marginal occurrence). As more data become
available and taxonomic issues are claried, it is possible
that some of these species may also prove to be threatened.
In the EU 28, six species (0.3%) are assessed as Regionally
Extinct, two are assessed as Extinct (0.1%). 65 species
(3.8%) are Critically Endangered, 150 species (8.7%)
are Endangered, and 183 species (10.6%) are Vulnerable.
A further 173 species (10%) are classied as Near
reatened. For 82 species (4.8%) in the EU 28 there
were insucient data to evaluate their risk of extinction
and so they were classied as Data Decient (Table 2,
Figure 4). Not Evaluated refers to species occurring at the
European level that did not occur within the EU Member
States (for example, only occurs in European Russia).
3.2 Status by taxonomic group
Table 3presents the status of European bryophyte species
organised by major group, split into mosses, liverworts
and hornworts. It is not considered useful to break it
down further into orders or families because the higher
classication of bryophytes is continually changing
with ongoing research. A stable consensus on the higher
classication of bryophytes will probably not be reached
for several years. ere are many more species of mosses
(1,327) than liverworts (461) and hornworts (8) in
Europe.
e percentages of species in dierent threat categories
are similar for mosses and liverworts. is similarity
may reect the fact that there is nothing particularly
distinctive about their ecology or distribution that
may inuence extinction risk. Instead, dierent life
strategies and ecologies are spread widely throughout
both liverworts and mosses, with plants in closely related
genera and families often having quite dierent ecological
requirements. e percentage gure for hornworts cannot
be directly compared with that for mosses or liverworts
because there are only eight species occurring in Europe.
Of these, only two species are considered endangered in
Europe.
Table 1. Proportion of threatened mosses, liverworts and hornworts in Europe and EU 28.
Europe
% species threatened
EU 28
% species threatened
Lower bound
(CR+EN+VU) / (assessed – EX) 21.4 23.1
Mid-point
(CR+EN+VU) / (assessed – EX – DD) 22.5 24.3
Upper bound
(CR+EN+VU+DD) / (assessed – EX) 26.6 27.9
12
Figure 4. IUCN Red List status of mosses, liverworts
and hornworts in Europe.
Figure 5. IUCN Red List status of mosses, liverworts
and hornworts in the EU 28.
Table 2. Summary of numbers of mosses, liverworts and hornworts within each Red List Category. Numbers of
endemic species are shown in brackets
IUCN Red List Categories No. species Europe
(no. endemic species)
No. species EU 28
(no. endemic species)
Extinct (EX) 2 (2) 2 (2)
Extinct in the Wild (EW) 0 (0) 0 (0)
Regionally Extinct (RE) 6 (0) 6 (0)
Critically Endangered (CR) 59 (18) 65 (15)
Endangered (EN) 143 (40) 150 (28)
Vulnerable (VU) 180 (36) 183 (25)
Near reatened (NT) 173 (33) 173 (22)
Least Concern (LC) 1140 (45) 1067 (13)
Data Decient (DD) 93 (17) 82 (11)
Total number of species analysed 1,796 (191) 1,728 (116)
Not Applicable (NA) 21 (0) 20 (0)
Not Evaluated (NE) - 69 (0)
Total number of species considered 1,817 (191) 1,817 (116)
Table 3. IUCN Red List status (at the European level) of mosses, liverworts and hornworts.
Total CR EN VU NT LC DD EX or
RE
Best estimate of %
threatened*
Mosses 1,327 43
(3.2%)
103
(7.8%)
137
(10.3%)
120
(9.0%)
853
(64.3%)
64
(4.8%) 7 (0.5%) 22.5
Liverworts 461 16
(3.5%)
38
(8.2%)
43
(9.3%)
51
(11.1%)
283
(61.4%)
29
(6.3%) 1 (0.2%) 22.5
Hornworts 8 0 2
(25.0%) 02
(25.0%)
4
(50.0%) 0 0 25.0
Total 1,796 59
(3.3%)
143
(8.0%)
180
(10.0%)
173
(9.6%)
1,140
(63.5%)
93
(5.2%) 8 (0.4%) 22.5
*e percentage of threatened species provides the mid-point gure as the best estimation of extinction risk. In addition, 21 NA species were not included in this table.
9.6
10
8
EN
VU
NT
LC
DD
RE
EX
3.3
0.3
0.1
63.5
5.3
CR
10
10.6
8.7
EN
VU
NT
LC
DD
RE
EX
3.8
0.3
0.1
61.7
4.8
CR
13
False dog-tooth Cynodontium fallax (Near threatened moss) © Tomas Hallingbäck
Polytrichum juniperinum (Least Concern moss) © Lars Hedenäs
14
3.3 Spatial distribution of species
3.3.1. Species richness
e geographic distribution of bryophyte richness in
Europe is shown in Figure 6 and is based on all native
and naturalised species (post 1500 AD) with extant and
possibly extant occurrence (1,796 species).
e areas with the highest species richness include central
Europe, namely mountainous areas in the Alps, and to
some degree in Scandinavia, Scotland, Wales, Pyrenees,
and Eastern Europe, including the Carpathians. Species
richness gradually declines towards the south and the
east of Europe. It is clear that mountainous areas score
most highly in terms of species richness. While there is
some overlap of species, each of these areas has its own
distinctive character, with the Scottish mountains, for
example, supporting a high diversity of Atlantic species,
in contrast to the Austrian Alps, where the ora is more
continental (see Box 2).
3.3.2. Endemic and near-endemic species
richness
In Figure 7, the richness of endemic European bryophyte
species is shown based on the presence of 184 species (the
analysis does not include species where their presence is
uncertain).
e incidence of endemic species is fairly constant
throughout most of Europe, with an increase in
hyperoceanic and mountainous areas, particularly
Macaronesia. Levels of endemism are low in bryophytes,
relative to vascular plants, although recent studies are
revealing an increasing number of previously unrecognised
endemic species (for example, Carter et al,. 2016, Patino
& Vanderpoorten, 2018). It is therefore more instructive
to look at areas where there is high diversity and large
numbers of ecologically specialised, disjunct and near-
endemic species. us, the hyperoceanic parts of Europe,
including Macaronesia, western Britain, Ireland, Norway,
France (Brittany) and north-western Spain, support rich
communities of oceanic species, few of which are endemic
(except in Macaronesia) but many are globally rare and
disjunct, elsewhere occurring only in widely-spaced but
climatically similar areas, such as Yunnan in China and
British Columbia in Canada (for example, Blockeel et al.,
2014). is is largely because of the very ecient dispersal
mechanisms in bryophytes, although some isolated and
disjunct populations may be relict. For example, the large
liverwort Anastrophyllum alpinum occurs in north-west
Figure 6. Species richness of European mosses, liverworts and hornworts.
15
Scotland, and elsewhere only in the Himalayas, Yunnan
and the Aleutian Islands. Similarly, southern Europe
supports a distinctive Mediterranean ora with many
species restricted to the Mediterranean basin, but not
necessarily endemic to Europe, occurring also in North
Africa, Turkey and adjacent countries (for example, Ros
et al., 2013). e tiny moss Acaulon fontiquerianum is a
rare species of southern Europe and the Canary Islands
that is also reported from Asiatic Turkey. ere are few
endemics in northern Scandinavia and Arctic Russia, but
the bryophyte ora is very distinctive and largely restricted
to the far north of Asia and North America, as well as
Europe: Drepanocladus arcticus is a strictly Arctic moss
conned to Svalbard, Arctic Russia (European and Asian)
and Arctic North America.
3.3.3. Distribution of threatened species
In Figure 8, the distribution of threatened bryophytes
in Europe is presented based on data for 374 threatened
species (the analysis does not include species where their
presence is uncertain).
Figure 8 displays the number of threatened species (CR,
EN, VU) per unit area (865 km
2
hexagon). As for overall
species richness (Figure 6), it shows a high number of
species in the Alps, especially in the eastern Alps, followed
by other mountainous areas, notably the Carpathians,
the eastern Pyrenees and the Scandinavian mountains.
is emphasises the importance of mountain habitats for
threatened bryophytes and their conservation. It may also
ag the impact of climate change on the mountainous
bryophyte ora, and pressure from land use change and
tourist developments in the high mountains. Furthermore,
many mountainous species are naturally rare and therefore
susceptible to stochastic events. Two regions notable for
their numbers of threatened species are an area located in
central Germany and Macaronesia (Figure 6). While the
high numbers in central Germany are dicult to explain,
the laurel forests of Macaronesian islands, which contain
many rare, threatened and endemic (Figure 7) bryophytes,
are under considerable threat from climate change,
wildres and forestry [see Box 2].
With the exception of these relatively restricted areas,
numbers of threatened species are fairly constant
throughout Europe. e map above (Figure 8) clearly
illustrates regions with a concentration of threatened
species, which deserve special attention for conservation
implementation. ey largely agree with the regions with
Figure 7. Distribution of endemic mosses, liverworts and hornworts in Europe.
16
Figure 8. Distribution of threatened (CR, EN, VU) mosses, liverworts and hornworts in Europe.
Figure 9. Distribution of Data Decient mosses, liverworts and hornworts in Europe.
17
Box 2 - Taking a closer look: Atlantic bryophytes
e extreme west of Europe, where the climate is warm and wet, neither too hot nor too cold, supports a rich
and varied selection of mosses and liverworts that are collectively referred to as oceanic or Atlantic. ese were
rst dened (in western Britain and Ireland) by the late Derek Ratclie in 1968, who identied the species that
are more or less conned to parts of Britain and Ireland which have more than 200 ‘wet days’ per year, with
wet days dened as days with over 1 mm of rain (Ratclie, 1968): in other words, the extreme west, where the
inuence of the Atlantic is at its strongest. e same suite of species extends to the Faroe Islands and, in less
abundance, to western Norway, western France (Brittany) and north-western Spain.
A lowland ravine in western Scotland may support upwards of 200 species in its sheltered, humid interior,
including many oceanic species. e Atlantic inuence in Scotland extends into the mountains, where a
community dened by Ratclie as the ‘mixed hepatic mat’ occurs in luxuriance in suitable north-east-facing
corries. is consists of large leafy liverworts such as species of Anastrophyllum, Bazzania, Herbertus, Plagiochila
and Scapania. Many of these species, as well as being strictly conned in Europe to the extreme west, are globally
rare and more or less threatened.
Herbertus borealis (Vulnerable liverwort) is endemic to Scotland © Michael Lüth
e oceanic ora is even better developed in Macaronesia, where the native laurel forest supports a very rich
assemblage of species, including a higher proportion of endemic species than is usual with bryophytes, especially
in Madeira. e subtropical forest is more or less constantly humid and warm and the trees are festooned with
mosses and liverworts, including tiny species of Acrobolbus and Lejeunea, as well as much larger Plagiochila and
Herbertus. ese small areas of forest are under great threat from climate change, wildres and developments
related to tourism.
Europe’s Atlantic bryophytes have clear anities with the tropical bryophyte ora of South America, and several
species once thought to be European endemics are now known to be identical with South American plants:
Plagiochila bifaria, for example, formerly known in Europe as P. killarniensis (Heinrichs et al., 1998). is is
probably the result of long distance dispersal of some species from South America to Europe via Macaronesia.
18
high general species richness, and partly with regions of
high levels of endemism (Figures 6 and 7).
However, it does not inform on relative regional threat
pressures. For this, the number of threatened species
would need to be represented as proportions of total
species number per region, a level of analysis that should
be done in future work. It is likely that lowland areas
which have experienced massive changes in land use due
to agricultural intensication and rural development since
the early 20th Century, still face a greater negative impact
than many mountainous areas; at least as long a climate
change has not yet have full eect on population decline.
3.3.4. Distribution of Data Decient species
In Figure 9, the distribution of Data Decient (DD)
species is presented based on data for 88 DD species (the
analysis does not include species where their presence is
uncertain). Some species are listed as DD because they
have been recently described and there is no information
to elucidate their trends, while others have been assessed
as DD due to taxonomic uncertainty and the diculty
to dierentiate between dierent species unless studied
genetically.
e incidence of DD species is often high in mountainous
areas, which could be attributed to the fact that they are
the most species-rich areas (Figure 6), but could also
be because they are usually more remote and dicult
to survey than the lowlands. ere are also more DD
species in relatively under-recorded parts of Europe, such
as Romania, than there are in well-recorded areas, such
as Britain and Ireland. e low number of DD species
throughout most of European Russia, may simply reect
the fact that much of it has low bryological diversity;
alternatively, it might indicate that this is an area which is
poorly surveyed and hence poorly known.
3.4 Major threats to moss, liverwort and
hornwort species in Europe
A comprehensive overview of the threats to bryophytes
in Europe is not possible, as some of the threats to the
species remain unknown. In total, it was possible to
identify threats for 1,099 species, often with multiple
threats listed for a species. Based on the best available
knowledge, 559 species are thought to have no current or
major threats, and for 159 species the threats are unknown
at present. reats to bryophytes are complex and often
dicult to categorise. ere are sometimes synergistic
eects between threats e.g., between climate change and
increased re frequency and it is hard to determine which
threat is the key driver impacting a bryophyte. In other
cases there may be several threats aecting an area e.g.,
climate change, increased re frequency and unregulated
planting of Eucalyptus and conifer plantations, etc., these
threats are all closely linked and it is hard to identify which
one is the key threat impacting a bryophyte species.
A summary of the major threats to threatened and not
threatened (DD, LC and NT) species is shown in Figure
10.
Natural system modications
Climate change and severe weather
Agriculture and aquaculture
Pollution
Human intrusions and disturbance
Residential and commercial development
Invasive and other problematic species, genes and deseases
Biological resource use
Energy production and mining
Transportation and service corridors
Geological events
Other options
0 100 200 300 400 500 600 700
reatened species
LC, NT and DD species
Figure 10. Major threats to all assessed mosses, liverworts and hornworts in Europe. Note: Species can be aected
by more than one threat.
19
e main threat to both mosses and liverworts is natural
system modication; 452 species of mosses and 180
species of liverworts are impacted by this driver of decline,
of which 144 and 52 species, respectively, are threatened.
For hornworts, the most prevalent threat is agriculture,
aecting seven species (of which two are threatened).
However, it should be noted that only eight hornwort
species were assessed in total, so this result should be
interpreted cautiously. Climate change ranked second in
the list of threats to bryophytes in Europe.
3.4.1 Natural system modications
A total of 234 species are aected by water management
and use, including 83 species assessed as CR, EN or
VU. is was considered the most common threat to
bryophytes across Europe, including species assessed
as threatened and species assessed as Near reatened
or Least Concern. is includes the abstraction of
ground and surface water for dierent uses, including
agricultural, commercial and domestic uses, and the
construction of dams. Species that are water-dependent,
such as Campylophyllum montanum and many of its
relatives, and those that tend to grow in sites targeted for
dam construction, such as Bryum blindii, are most at risk
from these threats.
Bogs and fens are among the most threatened habitats in
Europe (Janssen et al., 2016). In the lowlands, draining
wetlands has led to a substantial decline in many bog and
fen bryophyte species in central Europe (for example,
Sphagnum spp., Hamatocaulis vernicosus, Scorpidium
scorpioides, etc.), that are still quite common in the far north.
Remaining bryophyte-rich wetland sites, particularly in
central and southern Europe, face multiple threats, and all
require protection. In the uplands, construction of large-
scale dams and reservoirs has destroyed many rich sites,
and continues to threaten the survival of many species,
including those likely to be impacted by climate change
(for example, Andreaea crassinervia).
257 species appear to be at risk of an increase in re
frequency and/or intensity. Out of these, 94 species
are considered to be threatened. is threat is closely
associated with climate change, and becomes more serious
with a warmer and drier environment. e problem is
particularly serious in the laurel forests of Macaronesia,
where many rare and endemic species conned to these
forests are threatened by the increasing incidence of
wildres. ere is also a greater risk of wildres where
there has been large-scale planting of non-native Pinus
and Eucalyptus, as in much of southern Portugal and
northern Spain.
A total of 215 species are aected by a variety of other
modications to ecosystems, 75 of which have been
assessed as threatened.
3.4.2 Climate change
A total of 493 species are aected by climate change and
severe weather, of which 196 are threatened. Under this
Cheilolejeunea cedercreutzii (Endangered liverwort) © Tomas Hallingbäck
20
broad threat, 209 species are estimated to be or will be
aected by droughts, including 146 threatened species.
A total of 235 species are aected by habitat shifting and
alteration, including 109 threatened species. 163 species
are considered to be at risk from temperature extremes,
of which 78 species are threatened. With increasing
temperatures across Europe, as a result of climate change,
periods of droughts are already increasing (Vicente-
Serrano, 2014). e eects of climate change are often
unpredictable but the threat will only become more
prominent in the coming decades.
Some of the species most likely to be threatened by climate
change are those conned to wetlands. Already greatly
reduced, especially in central and southern Europe, due
to land-use changes including agricultural expansion
and intensication, drainage, pollution, construction
activities and invasive species, the remaining wetlands
are under extra pressure from desiccation caused by
climate change. Species of bryophytes that are found
at high elevations, and/or in northern environments,
are probably signicantly more prone to the impacts of
climate change than other species, as they have nowhere
else to go if temperatures increase signicantly. For
example, the survival of Herbertus sendtneri, a species of
the high Austrian Alps, is very doubtful if the extent and
duration of alpine snow-patches deteriorate signicantly.
On the other hand, at least the bryophytes of higher
elevations are usually less at risk from other factors; for
those that grow on lower mountains, climate change
is just one more threat to add to the other pressures
aecting them, such as land-use change.
e bryophytes of the laurel forests of Macaronesia
are also at great risk through climate change (Patiño
& Vanderpoorten 2018). e forests appear to be
drying out, wildres are becoming more frequent,
and projections show a signicantly increased risk of
extinction for many of the special species of this habitat
in the coming years, including endemic species such as
Cheilolejeunea cedercreutzii.
e bryophytes of southern Europe in general are also
at an increasingly higher risk of extinction as the climate
becomes warmer and drier. Already some areas have been
aected by desertication, and while many species have
strategies for avoiding or tolerating drought (for example
Gigaspermum mouretii, and many species in the family
Pottiaceae), even these species will be unable to survive
in conditions of more extreme desertication.
3.4.3 Agriculture and aquaculture
A total of 323 species are aected by agriculture and
aquaculture, of which 102 are threatened. Under this
broad threat, wood and pulp plantations aect 200
species of bryophyte in Europe, including 61 threatened
species. Dierent species are impacted by plantations
at dierent scales, but particularly by agro-industry
plantations. At this scale, 124 species are at risk, of which
26 are threatened.
Echinodium renauldii (Endangered moss) © César Garcia
21
Most plantations are on sites where there used to be
natural or semi-natural forest, so the main species
threatened by conversion of natural forest to plantation
woodland are those dependent on the long ecological
continuity provided by a stable, humid, natural forest.
Specialists of dead wood such as Scapania apiculata have
been particularly impacted, as amounts of deadwood are
often very low in managed forests.
Generally, land-use conversion practices (including the
intensication of agriculture and forestry) are considered
the most common threat to biodiversity in undisturbed
habitats (IPBES, 2018). ese have been designed
to increase the production of crops (for example, by
increased fertilizer and pesticide applications), livestock,
aquaculture, forest biomass, as well as urban development,
and are highly detrimental to bryophytes.
A total of 151 species are aected by livestock farming and
ranching, including 65 threatened species. is includes
grazing at three scales; nomadic, small-holders and agro-
industry. e majority of the species (95, of which 41
are threatened) are aected by small-holder grazing,
ranching or farming. Overgrazing, under-grazing, and
burning are all activities that may aect bryophytes.
One activity that is particularly associated with livestock
farming is the treatment of stock with ivermectins and
other chemicals to treat parasite infestations. One of the
unintended consequences of this is that it makes the dung
of these animals eectively sterile, which has knock-on
eects on the large numbers of organisms that depend on
animal dung for survival. ese include a unique suite of
mosses - the dung mosses - that grow only on the dung
(or sometime bones) of herbivores and which have an
intimate relationship with dung invertebrates for the
dispersal of their spores. Most of these species, some of
them among our most attractive bryophytes (Splachnum,
Tetraplodon, Aplodon, etc), have declined drastically in
recent years (Porley & Hodgetts, 2005).
3.4.4 Other threats to bryophytes
Residential and commercial development
A total of 185 species are aected by residential and
commercial development, of which 88 are threatened.
Under the umbrella of residential and commercial
development, a key threat to bryophytes in Europe was
identied as the development of areas for tourism and
recreation. 181 species in total are impacted by tourism
and recreation, including 99 threatened species. Tourism
encompasses many sorts of threat, including uncontrolled
building of hotels and other tourist facilities in rich
coastal or alpine habitats, water abstraction, disturbance
through increasing numbers of people, etc. ‘Urban
sprawl’ which occurs in order to accommodate the ever-
increasing human population, often replaces woodland,
species-rich grasslands and wetlands.
Pollution
A total of 240 species are aected by pollution, of which
66 are threatened. Under this broad threat, a total of 165
Dead wood is an important substrate for many specialised bryophytes; this old rotting tree trunk supports many species, including Scapania apiculata (Near reatened liverwort)
© Michael Lüth
22
species are aected by agricultural and forestry euents,
including 58 threatened species. is includes 97 species
(of which 34 of threatened) that are specically at risk from
nutrient loads, 29 species (of which ten are threatened)
that are specically at risk from herbicides and pesticides,
and 16 species (of which ten are threatened) that are
specically at risk from soil erosion and sedimentation.
Bryophytes are also considered to be aected by other
sorts of pollution in Europe. 78 species (of which 19 are
threatened) are impacted by air-borne pollutants, for
example acid rain and smog, and 32 species (of which
nine are threatened) are impacted by waste water, such as
run-o and sewage [see Box 3].
Invasive non-native/alien species/diseases
A total of 169 species are aected by invasive and other
problematic species, genes and diseases, of which 79
are threatened. Under this broad threat, 161 species are
aected by invasive alien species, including 79 threatened
bryophytes. Most of these problematic species are
unspecied, but 64 species (of which 40 are threatened) are
aected by known species. One of the most problematic
invasive plants for bryophytes in Europe is non-native
rhododendron (Rhododendron ponticum), which has
spread from gardens and now covers large areas of hillside
in oceanic areas, casting a deep shade and dropping very
acid leaf litter that prevents anything else from growing.
e eects of this invader on bryophytes are particularly
bad in areas of the UK and Ireland. However, it is
important to note that rhododendron (R. ponticum) is
a native and non-invasive plant in ravines in southern
Spain and Portugal; these rhododendron ravines are
threatened habitats and rich in bryophytes. e aquatic
environment is particularly sensitive to invasive species,
and plants such as Crassula helmsii are as much a threat
to aquatic bryophytes as they are to vascular plants. A
minority of species may be under threat from invasive
alien bryophytes: In north-western Europe, Orthodonium
gracile appears to be a poor competitor against the
invasive southern African species O. lineare.
Human intrusions and disturbance
A total of 204 species are aected by human intrusions
and disturbance, of which 92 are threatened. Under
this broad threat, human disturbance to areas where
bryophytes grow, specically for access to recreational
activities, is considered to impact 159 species, of which
81 are threatened. is includes intrusions relating to,
for example, erosion at popular tourist sites owing to the
sheer numbers of people; mountain summits in tourist
areas are particularly vulnerable in this respect. Some
coastal cli top paths in southern England with rare
mosses are becoming increasingly eutrophicated by dog
faeces, leading to loss of habitat. Hunting and shooting
for sport is not a direct threat to bryophytes, but when
important sites are managed primarily for these activities,
it can result in loss and degradation of habitat, as has
taken place on the grouse moors of Scotland, where large
areas are regularly burned to encourage the growth of
new heather (Calluna vulgaris) shoots, resulting not only
in a species-poor monoculture, but also destabilisation of
soils and increased erosion.
Additional threats
ere are many other threats to European bryophytes.
Biological resource use, which includes unintentional
impacts of shing, hunting and harvesting biological
resources, aects 152 species, of which 46 are threatened.
In addition, 116 species are threatened by industrial
activities, such as energy production and mining, including
31 threatened species. Under this threat classication,
most species (72, of which 12 are threatened) are at
risk from mining and quarrying, although renewable
energy production and oil and gas drilling also impact
some species in Europe. In some areas, such as central
Ireland, industrial-scale peat extraction for fuel has
damaged or destroyed many important bryophyte sites.
e remaining ones now receive statutory protection.
Land-based wind farms often cause considerable damage,
especially if sited on sensitive peaty substrates.
e establishment of transportation and service corridors,
such as roads and service lines, aects 69 species, of which
54 are threatened.
3.5 Population trends
Documenting the population trend of a species provides
key information when assessing its Red List status. As
part of this process, the whole population of each species
in Europe was assessed as declining, stable, increasing or
unknown.
Overall, 17.1% (307 species) of bryophyte species in
Europe are thought to be in decline, including 52.8% of
threatened species (162 species). e majority of species
(59.3%; 1,062 species) are considered to be stable,
including 8.3% of threatened species (88 species), and
1.9% (34 species) are increasing (Figure 11), all of which
23
are LC. However, 21.7% of species (389 species) have
unknown population trends, with 129 threatened species
(33.2%).
59%
Stable
17%
Decreasing
2%
Increasing
22%
Unknown
Figure 11. Population trends of European mosses,
liverworts and hornworts.
3.6 Gaps in knowledge
While there was not enough information to assign a Red
List Category to 93 species (hence considered as Data
Decient), the information collected was sucient to
identify the major knowledge gaps for bryophytes in
Europe (Figure 12).
Overall, the absence of, or the existence of few, data
on population size and distribution, as well as trends
are systematically highlighted as a knowledge gap for
bryophytes by the expert community assessing the
conservation status of these species. is pattern aects
both threatened and non-threatened taxa. Knowledge on
habitats trends and impact of threats is also still incipient
for the majority of these species, with particular regions
severely understudied (for example, Russia).
While this pattern can be partially justied by the fact
that some species have been recently described, and so
there is no information available on these parameters,
the reality is that monitoring eorts are becoming
increasingly dicult to sustain and to fund. is,
coupled with the absence of baseline data (for example,
historical data) on species numbers and distribution,
hamper a comprehensive understanding of the threats to
Figure 12. Research needs for European mosses, liverworts and hornworts. Note: Species can be included in more
than one category.
Research actions
Taxonomy
Life history and ecology
reats
Population size, distribution and trends
Population trends
Habitat trends
Species Action / Recovery Plan
Area-based Management Plan
5000 1000
Research
Monitoring
Conservation
Planing
reatened taxa
LC, NT and DD taxa
24
these species in Europe, and how these stressors interact.
Collecting information on these topics is paramount for
sound conservation planning and eective recovery of
threatened taxa, and will allow for more concrete messages
to be mainstreamed to the most impactful sectors.
e establishment of an expert network to facilitate
information exchange would certainly help address the
knowledge gaps identied for these species throughout
their European range; the experts brought together
through this project provide a good starting point to
expand this network. In any case, relevant conservation
and management measures should move ahead despite
any current data gaps, while also considering taxonomic
uncertainty where relevant.
Hymenoloma compactum (Data Decient moss) © Tomas Hallingbäck
25
Box 3 - Poisoned bryophytes: the impact of over-fertilization
One of the major threats to bryophytes is habitat modication through intensication of agricultural practices
and pollution (which may also come from agriculture). e greatest pollution threat in the 20th Century was
sulphur dioxide pollution through the widespread burning of dirty coal. Many bryophytes, particularly epiphytic
species, are very sensitive to sulphur dioxide (SO2) levels, and these plants, such as species of Orthotrichum and
Ulota, virtually disappeared from large areas of Europe. When legislation for clean air was introduced in the mid-
to late-20th Century, these plants gradually, and later rapidly, began to recolonise. Nowadays, trees throughout
most of Europe, even in areas formerly heavily impacted by SO2 pollution, are covered with Orthotrichum and
Ulota and other species.
Currently, the problem is nitrogen. Despite generally improving air quality in Europe, including reductions in
nitrogen emissions, there is an ever-increasing amount of nitrogen in the environment because of agricultural
practices and vehicle emissions. Locally, in agricultural areas, one can smell ammonia because of the enthusiastic
spreading of manure, and observe the homogenous green of ‘improved’ pasture devoid of wild owers or much
natural interest at all. Vehicle emissions are a big source of pollution by nitrogen compounds in densely populated
areas. However, the main cause is the worldwide increase in the very inecient use of articial nitrogen-rich
fertilisers in agriculture. In Europe, fertilisers are still spread over elds, but more than half the nitrogen does
not go into improved crop yields, it simply runs o into ditches, streams, rivers and ultimately the sea. Much
of it, via the nitrogen cycle, is returned to the land through precipitation. e consequences for bryophytes are
evident. All over Europe, even in remote upland areas, the natural species-rich bryophyte ora of streams is
being replaced with a monocultural slime of green algae; even in bogs, in some areas Sphagnum hummocks are
becoming overwhelmed by algal scum. On rock faces, a layer of green algae replaces the mosses and liverworts.
In open habitats, the nutrient-poor ‘bare ground’ habitat of so many threatened species is disappearing, being
overtaken by vigorous, nutrient-demanding grasses.
is is a worldwide problem that can only be addressed by worldwide solutions. Most agricultural land is
currently over-fertilised (Pearce, 2018), and so possible solutions include more targeted, ‘precision agriculture’,
distributing smaller amounts of nitrogen much more eciently, so that it goes to plant roots and does not run
o into the surrounding environment.
Urn bristle-moss Orthotrichum urnigerum (Vulnerable moss) © Michael Lüth
26
4. Conservation actions
4.1 Conservation of moss, liverwort and
hornwort species in Europe
e results of this Red List assessment indicate that 88.2%
of species (1,603 species, of which 319 are threatened)
were recorded in at least one protected area (including
national parks, Natura 2000 sites or nature reserves).
is is positive, as site protection is the most commonly
identied conservation action needed for European
bryophytes (Figure 13). e second most important
action is site/area management, and bryophytes are
often not considered in management plans. Additional
conservation measures proposed for European bryophytes
are shown below (Figure 13).
e nature conservation policy of the European Union
is based on two main pieces of EU legislation - the
1979 Birds Directive (Directive 79/409/EEC) and
the 1992 Habitats Directive (Directive 92/43/EEC;
jointly referred to as the Nature Directives). ere are
32 bryophyte species currently listed in Annex II of
the Habitats Directive, not all of which are endemic to
Europe. No species of bryophytes is listed under Annex
IV and only three genera are listed under Annex V of the
Habitats Directive. e Bern Convention, on the other
hand, is a binding international legal instrument that
aims to conserve wild ora and fauna and their natural
habitats and promote European cooperation towards
that objective. It covers all European countries and some
African states. In Appendix I of the Bern Convention
(Strictly Protected Flora Species), a total of 26 bryophyte
species are listed. Appendix 4 provides the full list of
bryophytes species listed under the Habitats Directive
and the Bern Convention, and the corresponding Red
List status as determined by this assessment. Of the
1,729 bryophyte species present in the EU 28, 7.5% are
endemic to the EU 28, highlighting the conservation
responsibility of the EU towards these species. Some
have made a remarkable recovery following listing under
the Habitats Directive and the Bern Convention, and
Figure 13. Main conservation actions needed identied for European mosses, liverworts and hornworts. Note:
More than one conservation action was assigned to each species.
Legislation
Awareness and communications
Formal education
Genome resource bank
Captive breeding/articial propagation
Habitat and natural process restoration
Invasive/problematic species controll
Site/area management
Resource and habitat protection
Site/are protection
0 100 200 300 400 500 600 700
Law and
policy
Education and
awareness
Spedies manage-
ment - Ex situ
conservation
Land / water
management
Land / water
protection
reatened taxa
LC, NT and DD taxa
27
targeted conservation actions (for example, Hamatocaulis
vernicosus - see Box 4).
One of the main tools to enhance and maintain
biodiversity in Europe is the Natura 2000 network of
protected areas, which currently consists of over 27,500
sites, it covers 18% of land territory but 27.5% land and
marine area (EC, 2018). Natura 2000 sites provide an
essential tool in conservation even if the sites were not
specically designated for the preservation of particular
bryophyte species, as indirectly the general protection of
habitats usually also benets the bryophytes. However,
it is sometimes necessary to target the ecological needs
of these small plants more directly, which becomes
challenging when conservation actions are usually
targeted at more charismatic and well-known organisms.
e well-supported agri-environmental schemes devised
to promote sustainable farming across Europe are a
good illustration of the limited eects on bryophytes
of such widespread, untargeted conservation measures,
particularly for rare bryophyte species (Valentini et al.,
2016). Actions better tailored to promote bryophyte
conservation include, for example, rotational set-aside
and retention of winter stubbles in cereal, rape and
linseed crops (Bosanquet, 2003; Bisang et al., 2009).
Many threatened bryophytes occur in protected areas,
and depend, like other groups, on the conservation of
multi-scale areas of semi-natural habitat. However, many
species tend to grow in ‘micro-habitats’ in non-protected
areas. is means that the sympathetic management
of the wider countryside is particularly important for
bryophytes. For example, the suite of species which have
their main habitat in arable elds, such as the threatened
hornwort Anthoceros neesii, are entirely dependent
for their survival on overwintering stubble elds, so
wider agricultural policy needs to promote agricultural
practices that favour this habitat. Similarly, a certain
amount of dead wood needs to be left in situ in managed
forestry plantations, as well as in old-growth forest, in
order to provide substrate for the many bryophytes that
specialise in this habitat. While the Habitats Directive
Soft brook-moss Platyhypnum molle (Vulnerable moss) © Tomas Hallingbäck
28
and the Natura 2000 network are of vital importance,
there needs to be much more coordination between
dierent elements of policy, so that, to choose the most
obvious example, policies contained within the Common
Agricultural Policy (CAP) do not work against those in
the Habitats Directive. Measures within other national
or international policies, including Agri-Environmental
Schemes that provide payments to farmers who subscribe,
on a voluntary basis, to environmental commitments
related to the preservation of the environment, need to
be aligned with statutory protection for species to ensure
that eorts to protect these species are synergistic and not
in vain.
European countries and EU Member States are signatories
to a number of important conventions aimed at conserving
biodiversity, including the 1979 Bern Convention on the
Conservation of European Wildlife and Natural Habitats,
and the 1992 Convention on Biological Diversity (CBD).
rough the CBD, the Strategic Plan 2011–2020 was
established, which includes 20 targets (Aichi Targets) that
guide the implementation of the CBD and all the other
biodiversity conventions. In particular, Target 12 focuses
on preventing the extinction of known threatened species
and improving their status (CBD, 2011). e outcomes
of this Red List project certainly help to measure the
progress made towards meeting these targets, and the
current results suggest that, for bryophytes, Europe is not
on track to meet these targets.
e Global Strategy for Plant Conservation (GSPC)
was adopted by the CBD at the 2002 Conference of the
Parties and updated at the 10th Conference of the Parties.
In order to coordinate the implementation of the GSPC
at the regional level, the European Strategy for Plant
Conservation (ESPC) was adopted. In particular, Target
2 (calling for an assessment of the conservation status
of plant species), Target 5 (through the identication of
Important Plant Areas), Target 7 (in situ conservation),
Target 8 (ex situ conservation), Target 12 (preventing the
extinction of known threatened species and improving
their status), Target 13 (sustainable practices associated
with plant use) and Target 14 (awareness raising) (CBD,
2011) are relevant for the conservation of bryophytes.
European countries across the continent endorsed the
Pan-European 2020 Strategy for Biodiversity (UNEP,
2011), which re-focuses eorts to prevent further loss of
biodiversity in the region. It also provides a European
mechanism for supporting the implementation of
the global Strategic Plan for Biodiversity. No native
European bryophyte species are listed in the Appendices
of the Convention on International Trade in Endangered
Species of Wild Fauna and Flora (CITES). e EU
Water Framework Directive, adopted in 2000 and aimed
at protecting European waters, can also be relevant for
aquatic and water-dependant bryophyte species. A good
ecological status of surface waters, as promoted by the
Grey-cushioned grimmia Grimmia pulvinata (Least Concern moss) © Michael Lüth
29
Directive, has positive eects on ecosystem function
(Janauer et al., 2015).
Plant habitat conservation eorts have in part been
focused through the identication of Important Plant
Areas (IPAs). IPAs are internationally signicant sites
for wild plants and threatened habitats. Identied at a
national level, they provide a framework for implementing
Target 5 of the CBD GSPC, and are a tool for targeting
conservation actions on wild plants and in situ habitat
protection. IPAs contain over 700 of the most threatened
species in Europe and include millions of hectares of
the most threatened habitats. At least 1,770 IPAs have
been identied in 16 European countries (Anderson &
Radford, 2010). A rst attempt was made to identify
Important Bryophyte Areas in Europe during the
production of the rst Red List (European Committee
for the Conservation of Bryophytes, 1995), and it is
anticipated that the current Red List will facilitate an
update of this initiative. ese exercises are incredibly
valuable to ensure species protection and a stepping stone
to promote nature conservation in Europe.
e EU has committed to a long-term (2050) vision
and mid-term headline target for biodiversity, which
is “To halt the loss of biodiversity and the degradation of
ecosystem services in the EU by 2020 and restore them in
so far as possible, while stepping up the EU contribution to
averting global biodiversity loss.” is target underpins the
EU Biodiversity Strategy 2011–2020. e establishment
of these policy instruments indicates the high political
commitment to biodiversity and the need to monitor
the status of biodiversity and to assess progress towards
meeting conservation objectives and targets. Measuring
whether policy targets have been met is only possible by
establishing comprehensive monitoring programmes that
allow the gathering of the necessary data for a reliable re-
assessment in the coming years. e results of the present
Red List assessment indicate that, for the bryophytes,
Europe is currently not on track to meet these targets.
For the latter to happen, immediate conservation action
for species with a high extinction risk is needed.
Most European countries have developed specic actions
at the national or regional level in order to enhance
bryophyte populations. National Red Lists or Red
Data Books of bryophyte species are available for the
following countries: Austria, Bulgaria, Czechia, Estonia,
Finland, Germany, Great Britain (excluding Northern
Ireland), Hungary, Ireland (including Northern Ireland),
Tayloria rudolphiana (Near reatened moss) © Norbert Schnyder
30
Italy (including Sardinia), Luxembourg, Montenegro,
Netherlands, Norway, Poland, Portugal (including
Madeira), Romania, Serbia, Slovakia, Slovenia, Spain
and the Canary Islands, Sweden, and Switzerland.
However, there are some countries in which no national
Red List has been developed (for example, France). It is
also noteworthy that some national Red Lists are outdated
and should be maintained and updated in order to remain
relevant. In addition, several countries have developed
management or action plans for several species, and have
legislation in place to protect certain species legally (for
example, Schedule 8 of the Wildlife & Countryside Act
1981 in the UK). Some examples of successful action
plans include Tayloria rudolphiana, epiphytic species
of central Europe for which several studies have been
undertaken to count the sites or individuals of the plants
(Hofmann et al., 2006; Hofmann et al., 2016; Müller,
2016; Kiebacher et al., 2018) and attempts have been
made to protect the host trees and to increase awareness
for this species. LIFE projects have been undertaken
at the European level to enhance the status of certain
habitats and species, some of which have focused on
specic species bryophytes, or produced management
plans as a result of these projects.
4.2 Red List versus priority for conservation
action
Assessing the extinction risk and setting conservation
priorities are related but distinct processes. e purpose of
the IUCN Red List assessment is to produce an estimate
of the likelihood of extinction of a species. On the other
hand, setting conservation priorities also takes into
account other factors such as ecological, phylogenetic,
historical, economical or cultural preferences for some
taxa over others. Also, the probability of success of
conservation actions, availability of funds or personnel,
cost-eectiveness and legal frameworks for the
conservation of threatened taxa is taken into account.
In the context of regional risk assessments, a number of
additional pieces of information are valuable for setting
conservation priorities. For example, it is important to
consider not only conditions within the region, but also
the Red List status of the taxon from a global perspective
and the proportion of the global population that occurs
within the region. e decision on how these three
variables, and the other factors, are used for establishing
conservation priorities is a matter for the regional
authorities to determine, taking into account the Red
List status of the species of concern.
31
Box 4 - Conservation works: bryophytes bounce back
For bryophytes, as for other organisms, conservation works. Bogs that have been damaged by peat-cutting or
drainage can be restored or re-instated by blocking drains; degraded forest can be restored (eventually) merely by
non-intervention; damaged wetlands can have a new lease of life through proper management. Giving species
legal protection can be very eective. For example, Slender green feather-moss (Hamatocaulis vernicosus) was
one of a small handful of species placed on Appendix I of the Bern Convention in the early 1990s. Along with
most of the other Bern species, it was also included on Annex II of the EU Habitats Directive soon afterwards.
is means that the signatory countries to these conventions have an obligation to protect it under the Natura
2000 network, with sites designated and managed for its protection. e results of this have been dramatic,
with sites established for Hamatocaulis all over Europe, or at least within the EU, with several non-EU countries
following suit. Because it is a key species of mineral-rich, mesotrophic mires, fens and ushes, this has meant that
many important and threatened wetland sites that might otherwise have been destroyed now receive statutory
protection. In other words, the conservation benets of placing this moss on the protected species list extend
much further than merely protecting the moss itself: whole habitats have been saved.
Furthermore, including Hamatocaulis on these international conventions has resulted in a massively increased
programme of research to nd out more about its distribution, abundance, ecology and conservation
requirements. Recent research in Sweden has even found that what we call H. vernicosus actually comprises two
cryptic species (genetically dierent but apparently morphologically identical), both of which occur in protected
areas (Hedenäs, 2018). Paradoxically, an increase in survey and recording eort targeted at H. vernicosus means
that it now seems to be more common than was once thought. is is not the case: we simply now know more
about it, and the increase in records in recent years is entirely due to that increase in recording eort. If we
had more baseline data going back through the decades it would certainly show a decline because of habitat
destruction through drainage and other anthropogenic factors.
Slender green feather-moss Hamatocaulis vernicosus (Vulnerable moss) © Michael Lüth
32
5. Recommendations
5.1 Recommended actions
Currently, 22.5% of bryophytes are threatened at the
European level. e most important threats to bryophytes
in Europe come from natural systems modications (i.e.,
habitat destruction and degradation), climate change,
and current agricultural practices. Hence, improving
the conservation status of bryophytes, and preventing
current and future declines in Europe, requires increasing
eorts and commitments from various parties, from the
EU to regional assemblies, and from statutory bodies
to conservation charities. Perhaps most importantly,
measures for bryophyte conservation (and indeed for
nature conservation generally) need to be integrated
into regular planning and land management procedures
and practices. Below, a series of recommendations
are proposed to strengthen the long-term survival of
European bryophytes:
Policy measures
• Use the European Red List as the scientic basis to
inform regional/national lists of rare and threatened
species and to identify priorities for conservation
action in addition to the requirements of the Habitats
Directive, thereby highlighting the conservation status
of bryophytes at the regional/local level.
• Use the European Red List to support the integration
of conservation policy with the Common Agricultural
Policy (CAP) and other national and international
policies. For example, CAP Strategic Plans should
include biodiversity recovery commitments that could
anticipate, among others, the creation of Important
Bryophyte Areas. An increased involvement of
national environmental agencies in the preparation
of these strategic plans, and more broadly in ongoing
discussions on the Future CAP Green Architecture,
would likely also ensure the design of conservation
measures better tailored to conserve bryophytes in
agricultural landscapes.
• Update the European Red List every decade to ensure
that the data remains current and relevant.
• Develop Key Biodiversity Areas for bryophytes in
Europe with a view to ensuring adequate site-based
protection for bryophytes.
Research and monitoring
• Use the European Red List as a basis for future targeted
eldwork on possibly extinct and understudied species.
• Establish a monitoring programme for targeted
species (for example, threatened species and/or arable
bryophytes).
• Use the European Red List to obtain funding for
research into the biology and ecology of key targeted
species.
Action on the ground
• Use the European Red List as evidence to support
multi-scale conservation initiatives, including
designation of protected areas, reform of agricultural
practices and land management, habitat restoration
and rewilding, and pollution reduction measures.
• Use the European Red List as a tool to target species
that would benet the most from the widespread
implementation of the solutions oered by the 1991
Nitrates Directive (Council Directive 91/676/EEC),
including the application of correct amounts of
nutrients for each crop, only in periods of crop growth
under suitable climatic conditions and never during
periods of heavy rainfall or on frozen ground, and the
creation of buer zones to protect waters from run-o
from the application of fertilizers.
Ex situ conservation
• Undertake ex situ conservation of species of
conservation concern in botanic gardens and spore
and gene banks, with a view to reintroduction where
appropriate.
Awareness raising
Mosses and liverworts are small and do not impinge
very much on the public consciousness, except as things
to remove from the lawn or the roof. As an integral and
important part of the natural world, they deserve better.
ere are now many attractive publications and websites
that present bryophytes as beautiful and useful, and these
should receive more publicity and promotion; for example,
Sphagnum mosses - e Stars of European Mires (Laine et
33
al., 2018), Robert Muma’s beautiful moss paintings and
sketches
(http://worldofmosses.com/paintings/index.html)
,
Michael Lüth’s amazing photographic collection (http://
www.milueth.de/Moose/index.htm), to name but a few).
Many nature reserves where bryophytes are important
now have information boards and other material to
promote bryophytes, and this should continue to be
prioritised wherever appropriate.
In particular, this Red List should be used to publicise
bryophytes and to obtain funding for future conservation
work. For example, LIFE+ Nature and Biodiversity
2
provides targeted funding for species conservation actions,
supporting projects aimed at conserving threatened
species listed in the annexes of the EU Habitats Directive,
Birds Directive and the European Red List.
5.2 Application of project outputs
e European Red List of mosses, liverworts and hornworts
is part of a wider initiative aimed at assessing the status of
all European species. It provides key resources for decision
makers, policy makers, resource managers, environmental
planners, NGOs and the concerned public by compiling
large amounts of data on the population, ecology, habitats,
threats and recommended conservation actions for each
bryophyte species. Red List assessments are intended to
be policy-relevant and can be used to inform conservation
planning and priority setting processes. However, they
are not intended to be policy-prescriptive and are not in
2 http://ec.europa.eu/environment/life/project/Projects/index.
cfm?fuseaction=home.getProjects&strandID=2
themselves a system for setting biodiversity conservation
priorities. ese data are freely available on the IUCN
Red List website (https://www.iucnredlist.org/regions/
europe), on the European Commission’s website (http://
ec.europa.eu/environment/nature/conservation/species/
redlist) and through paper publications (see the list of
European Red Lists published at the end of this report).
Red Lists are a dynamic tool that will evolve with time as
species are re-assessed according to new information or
situations. ey are aimed at stimulating and supporting
research, monitoring and conservation action at local,
regional and international levels, especially for threatened,
Near reatened and Data Decient species.
Each species assessment lists the major threats aecting
the specic bryophyte species and conservation actions
that are in place or recommended. is is useful to inform
the application of conservation actions for each species.
e outputs of this project can be applied to inform
policies and to identify priority sites for biodiversity and
priority species to include in research and monitoring
programmes.
5.3 Future work
rough the strong collaboration established between the
ECCB and the IUCN SSC Bryophyte Specialist Group
during this project, a network of European and national
bryophyte experts, and their extensive knowledge and
expertise, were mobilised that will persist long after the
project ends and will be instrumental in dening priorities
Plagiomnium confertidens (Vulnerable moss) © Elvira Baisheva
34
for bryophyte conservation in Europe. e project has
beneted greatly from the work and information held
by additional relevant organisations and stakeholders,
such as national bryophyte societies, university research
programmes and statutory and voluntary conservation
bodies. e wealth of knowledge and data compiled
during the elaboration of this European Red List will be
invaluable to expand research eorts on bryophytes at the
European level, ultimately beneting their conservation.
One aspect worth noting is that the assessment of the
European endemics can be transcribed directly into the
corresponding global Red List.
rough the process of compiling data for the European
Red List, a number of knowledge gaps have been
identied. Across Europe there are signicant geographic,
geopolitical and taxonomic biases in the quality of data
available on the distribution and status of species, and
these are the aspects that a unied knowledge network
will need to overcome to advance bryophyte conservation
in the region.
ere is a clear need for drawing together information from
all data compilation initiatives, under way or planned, and
for a wider European bryophyte conservation action plan
to be explored, developed and progressed. It is hoped that
by presenting this assessment, local, national, regional
and international research will be stimulated to provide
new data and to improve on the quality of the current
available data.
Key challenges for the future are to improve monitoring,
research and data quality and dissemination so that the
information and analyses presented here can be updated
and improved. is will contribute to recommend
conservation actions based on a solid scientic basis. e
further dissemination of this information to concerned
European citizens will also lead to progressive policies at
various jurisdictional levels that promote conservation.
ere is also a need for education, both of the general
public and those involved in nature conservation, to
raise awareness and to take bryophytes into account in
conservation initiatives.
If the bryophyte assessments are periodically updated,
they will enable the changing status of these species
to be tracked over time via the production of a Red
List Index (Butchart et al., 2004; 2005; 2006; 2007).
To date, this indicator has been produced for birds,
mammals, amphibians and reptiles at the European
level and has been adopted as one of the headline
biodiversity indicators to monitor progress towards
halting biodiversity loss in Europe by 2020 (EEA, 2007).
e development of such an index will be important to
evaluate progress towards meeting Target 6 of the EU
Biodiversity Strategy and for discussions shaping the
Post-2020 Biodiversity Framework in order for Europe
to step up its contribution to averting global biodiversity
loss, and Aichi Target 12 of the CBD, which focuses on
preventing the extinction of known threatened species
and improving their status.
Sphagnum arcticum (Near reatened moss) © Michael Lüth
35
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European Red List of grasshoppers, crickets and bush-crickets.
Luxembourg: Publications Oce of the European Union.
doi:10.2779/60944
Hodgetts, N.G. (2015). Checklist and country status of European
bryophytes – towards a new Red List for Europe. Irish Wildlife
Manuals, No. 84. National Parks & Wildlife Service,
Department of Arts, Heritage & the Gaeltacht, Ireland.
Hofmann, H., Senn-Irlet, B. and Stofer, S. (2006).
Prioritätensetzung für Pilze, Flechten und Moose im Kanton
Bern. Naturschutzinspektorat des Kantons Bern, Bern.
Hofmann, H., Schnyder, N. and Kiebacher, T. (2016).
Rudolphis Trompetenmoos – Nachsuche im Kanton Bern.
Abteilung Naturförderung des Kantons Bern (ANF).
International Union for Conservation of Nature (IUCN)
(2011). ‘Guidelines for reporting on proportion threatened.
Version 1.0.’ In: IUCN (2011). Guidelines for appropriate
uses of IUCN Red List data. Incorporating the guidelines
for reporting on proportion threatened and the guidelines on
scientic collecting of threatened species. Version 2. Adopted
by the IUCN Red List Committee and IUCN SSC Steering
Committee. Downloadable from: http://www.iucnredlist.
org/documents/RL_Guidelines_Data_Use.pdf
IUCN (2012a). IUCN Red List categories and criteria: Version
3.1. Second edition. Gland, Switzerland and Cambridge,
UK: IUCN.
IUCN (2012b). Guidelines for application of IUCN Red List
criteria at regional and national levels. Version 4.0. IUCN
Species Survival Commission. Gland: IUCN.
IUCN (2014). Guidelines for using the IUCN Red List categories
and criteria. Version 11. Prepared by the Standards
and Petitions Subcommittee. Gland, Switzerland and
Cambridge, UK: IUCN. Available at: http://www.
iucnredlist.org/documents/RedListGuidelines.pdf.
IUCN (2015). European species under threat. Overview of
European Red Lists results. Downloadable from: https://
www.iucn.org/downloads/red_list_overview_new_1.pdf
Janauer, G.A., Albrecht, J. and Stratmann, L. (2015). ‘Synergies
and conicts between Water Framework Directive and
Nature 2000: legal requirements technical guidance and
experience from practice’. In: Ignar, S. and Grygoruk,
M. (eds.), ‘Wetlands and Water Framework Directive.
Protection, management and climate change’. GeoPlanet:
Earth and Planetary Sciences. Springer Open, ISSN 2190-
5193, pp. 9–29.
Kiebacher, T., Bergamini, A., Scheidegger, C. and Bürgi, M.
(2018). Bergahornweiden im Alpenraum: Kulturgeschichte,
Biodiversität und Rudolphis Trompetenmoos. Haupt.
Laine, J., Flatberg, K.I., Harju, P., Timonen, T., Minkkinen,
K., Laine, A., Tuittila, E.-S. and Vasander, H. (2018).
Sphagnum mosses - the stars of European mires. Helsinki,
University of Helsinki.
Meinunger, L. and Schröder, W. (2007). Verbreitungsatlas der
Moose Deutschlands - Band 2. Oliver Dürhammer für die
Regensburgische Botanische Gesellschaft, Regensburg.
Morris, J.L., Puttick, M.N., Clark, J.W., Edwards, D., Kenrick,
P., Pressel, S., Wellman, C.H., Yang, Z., Schneider, H. and
Donoghue, P.C.J. (2018). ‘e timescale of early land plant
evolution’. Proceedings of the National Academy of Sciences of
the United States of America 115(10): E2274-E2283. Doi:
10.1073/pnas.1719588115
Müller, N. (2016). Überraschend üppige Population von
Tayloria rudolphiana auf dem Grüscher Älpeli. Meylania
58: 14-18.
Patiño, J. and Vanderpoorten, A. (2018). ‘Bryophyte
Biogeography’. Critical Reviews in Plant Sciences 37:2-3,
175-209, doi: 10.1080/07352689.2018.1482444
37
Pearce, F. (2018). ‘Can the World Find Solutions to the
Nitrogen Pollution Crisis?’. Yale Environment 360.
Available at: https://e360.yale.edu/features/can-the-world-
nd-solutions-to-the-nitrogen-pollution-crisis
Pedroli, G.B.M. and Meiner, A. (2017). Landscapes in
transition. No. 10/2017. European Environment Agency
(EEA).
Porley, R. and Hodgetts, N. (2005). Mosses and Liverworts.
New Naturalist No. 97. HarperCollins, London.
Puttick, M.N., Morris, J.L., Williams, T.A., Cox, C.J.,
Edwards, D., Kenrick, P., Pressel, S., Wellman, C.H.,
Schneider, H., Pisani, D. and Donoghue, P.C.J. (2018).
‘e Interrelationships of Land Plants and the Nature of
the Ancestral Embryophyte’. Current Biology 28, 733–745.
doi: 10.1016/j.cub.2018.01.063
Ratclie, D.A. (1968). ‘An ecological account of Atlantic
bryophytes in the British isles’. New Phytologist 67: 365-
439.
Ros, R.M., Mazimpaka, V., Abou-Salama, U., Ale, M.,
Blockeel, T.L., Brugués, M., Cros, R.M., Dia, M.G.,
Dirkse, G.M., Draper, I., El-Saadawi, W., Erdağ, A.,
Ganeva, A., Gabriel, R., González-Mancebo, J.M., Granger,
C., Herrnstadt, I., Hugonnot, V., Khalil, K., Kürschner,
H., Losada-Lima, A., Luís, L., Mifsud, S., Privitera, M.,
Puglisi, M., Sabovljević, M., Sérgio, C., Shabbara, H.M.,
Sim-Sim, M., Sotiaux, A., Tacchi, R., Vanderpoorten, A.
and Werner, O. (2013). ‘Mosses of the Mediterranean, an
annotated checklist’. Cryptogamie, Bryologie 34: 99-283.
doi: 10.7872/cryb.v34.iss2.2013.99
Söderström, L., Urmi, E. and Váňa, J. (2007). ‘e distribution
of Hepaticae and Anthocerotae in Europe and Macaronesia
– Update 1–427’. Cryptogamie, Bryologie 28: 299–350.
Stark, L.R., Greenwood, J.L. and Brinda, J.C. (2016).
‘Desiccated Syntrichia ruralis shoots regenerate after 20
years in the herbarium’. Journal of Bryology 39(1): 85-93.
doi: 10.1080/03736687.2016.1176307
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Revision. United Nations, Department of Economic and
Social Aairs. Available online from: http://esa.un.org/
unpd/wpp/ExcelData/population.htm. Accessed on 8
September 2016.
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With a focus on: Cooperation for the conservation and
sustainable use of PanEuropean biodiversity and the
coordinated national implementation of biodiversity-related
Multilateral Environmental Agreements (MEAs). Nairobi:
United Nations Environment Programme.
Valentini, M., Bisang, I., Jacot, K. and Bergamini, A. (2016).
‘Do Bryophytes Prot from Agri-environmental Schemes?
A Comparison of Dierent “Ecological Focus Area” in the
Swiss Lowlands’. Conference paper. 9th ConferenCe of
European Committee for Conservation of Bryophytes. 26-
29. April 2016. Bečići, Montenegro.
Vicente-Serrano, S.M., Lopez-Moreno, J.I., Beguería,
S., Lorenzo-Lacruz, J., Sanchez-Lorenzo, A., García-
Ruiz, J.M., Azorin-Molina, C., Morán-Tejeda, E.,
Revuelto, J. and Trigo, R. (2014). ‘Evidence of increasing
drought severity caused by temperature rise in southern
Europe’. Environmental Research Letters 9(4): 1-9. Doi:
10.1088/1748-9326/9/4/044001
Villareal, J.C., Cargill, D.C., Hagborg, A., Söderström, S. and
Renzaglia, K.S. (2010). ‘A synthesis of hornwort diversity:
patterns, causes and future work’. Phytotaxa 9: 150-166.
Doi: 10.11646/phytotaxa.9.1.8
39
Appendix 1. List of lead assessors
by geographical region
• Central Europe: Norbert Schnyder and Christian Schröck
• Eastern Europe: Nadya Konstantinova and Elvira Baisheva
• Macaronesia: Manuela Sim Sim
• Northern Europe: Tomas Hallingbäck
• Northwestern Europe: Nick Hodgetts
• Southern Europe: Patrizia Campisi and Annalena Cogoni
• Southeastern Europe: Marko Sabovljevic
• Southwestern Europe: Cecilia Sérgio
Appendix 2. Example of species
summary and distribution map
e Red List assessment of Orthotrichum urnigerum on the following pages provides an example of the information
that has been compiled for all the European bryophyte species, including a distribution map. You can search for and
download all the assessments and distribution maps from the European Red List website and data portal available
online at http://ec.europa.eu/environment/nature/conservation/species/redlist/ and https://www.iucnredlist.org/
regions/europe.
40
2
Encalypta mutica - I. Hagen
PLANTAE - BRYOPHYTA - BRYOPSIDA - ENCALYPTALES - ENCALYPTACEAE - Encalypta - mutica
Common Names: Trubbklockmossa (Swedish)
Synonyms: No Synonyms
Red List Status
VU - Vulnerable, C2a(i) (IUCN version 3.1)
Red List Assessment
Assessment Information
Reviewed?
Date of Review:
Status:
Reasons for Rejection:
Improvements Needed:
true
2017-07-26
Passed
-
-
Assessor(s): Hodgetts, N., Blockeel, T., Konstantinova, N., Lönnell, N., Papp, B., Schnyder, N., Schröck, C., Sergio, C. &
Untereiner, A.
Reviewer(s): Wilbraham, J. & Cálix, M.
Assessment Rationale
European regional assessment: Vulnerable (VU)
EU 28 regional assessment: Vulnerable (VU)
Encalypta mutica is an essentially Arctic species that for a long time was thought to be a Scandinavian endemic, but it is
now also known from Estonia and Arctic European Russia. This rare species is assessed as Vulnerable since it is estiated that
there are fewer than 10,000 individual-equivalents in Europe and in the EU 28, and that each subpopulation has fewer than
1,000 individual-equivalents. One individual-equivalent (i.e., mature individual) is considered to be one square meter on
which the species grows. In addition, the current population trend is decreasing slightly.
Although the threats to this species appear to be largely unknown, it is certainly threatened, as it is both rare and
apparently declining. It appears that it may be threatened by a lack of grazing in some localities. In the south part of
Scandinavia, it is affected by the overgrowing of higher vegetation. Three localities were taken under monitoring in Estonia,
where either grazing or simulation of grazing is needed in order to re-create and preserve suitable open soil patches for this
species. Similar management can be recommended elsewhere. Research into threats is also recommended.
Distribution
Geographic Range
Encalypta mutica is an essentially Arctic species that for a long time was thought to be a Scandinavian endemic, but it is
now also known from Estonia and Arctic European Russia. The record from the Czech Republic was excluded in the 2003
edition of the Czech Red List (Kučera and Váňa 2003), but is considered at best uncertain in this assessment. Old records
from Romania are probably errors (S. Stefanut pers. comm. 2016). There are also records from the lowlands of Ukraine but,
without further details or specimens, these have to be regarded with scepticism. Elsewhere it occurs in Siberia, Greenland
and northern North America. This species' area of occupancy (AOO) is estimated at 452 km², and its extent of occurrence
(EOO) at ca 2.9 million km².
41
3
Elevation / Depth / Depth Zones
Elevation Lower Limit (in metres above sea level): 0
Elevation Upper Limit (in metres above sea level): 900
Map Status
Map
Status
How the map was created,
including data
sources/methods used:
Please state
reason for map
not available:
Data
Sensitive?
Justification
Geographic range
this applies to:
Date
restriction
imposed:
Done
-
-
-
-
-
-
Biogeographic Realms
Biogeographic Realm: Palearctic
Occurrence
Countries of Occurrence
Country
Presence
Origin
Formerly Bred
Seasonality
Czechia
Presence Uncertain
Native
-
Resident
Estonia
Extant
Native
-
Resident
Finland
Extant
Native
-
Resident
Norway
Extant
Native
-
Resident
Romania
Presence Uncertain
Native
-
Resident
Russian Federation
Extant
Native
-
Resident
Russian Federation -> European Russia
Extant
Native
-
Resident
Russian Federation -> European Russia -> North European Russia
Extant
Native
-
Resident
Svalbard and Jan Mayen
Extant
Native
-
Resident
Sweden
Extant
Native
-
Resident
Ukraine
Presence Uncertain
Native
-
Resident
Ukraine -> Ukraine (main part)
Presence Uncertain
Native
-
Resident
Population
This species may have disappeared from some of its Norwegian localities, including the type locality near Trondheim
(Hallingbäck et al. 2006). It seems to be threatened and declining in Estonia, where, in 2006, it was not found at one
previously known locality (Vellak & Ingerpuu 2012). In Murmansk there is only one locality (where it is rare), and a single
locality in Karelia and in the Polar Urals. The species is rather overlooked in the mountains where it is stable, although in
the lowlands in can be declining in limestone quarries. The overall current population trend is considered to be decreasing
slightly. The population is not severely fragmented. It is estimated that there are fewer than 10,000 individual-equivalents
in Europe and in the EU 28, and that each subpopulation has fewer than 1,000 individual-equivalents. One individual-
equivalent (i.e., mature individual) is considered to be one square meter on which the species grows.
42
4
Habitats and Ecology
This essentially Arctic species grows on bare calcareous, and periodically wet, soil, typically in very sun-exposed situations,
for example on alvar heaths, in limestone quarries and on gravel produced by weathering at the base of south-facing,
calcium-rich alpine slopes. Associates include Ditrichum flexicaule, Encalypta vulgaris, Myurella julacea and Weissia
controversa. The altitudinal range is from sea level up to 900 m Asl.
IUCN Habitats Classification Scheme
Habitat
Season
Suitability
Major Importance?
4.2. Grassland -> Grassland - Subarctic
Resident
Suitable
Yes
6. Rocky areas (eg. inland cliffs, mountain peaks)
Resident
Suitable
Yes
Systems
System: Terrestrial
Use and Trade
General Use and Trade Information
This species is not utilised or traded.
Threats
Although the threats to this species appear to be largely unknown, it is certainly threatened, as it is both rare and
apparently declining. It appears that it may be threatened by a lack of grazing in some localities. In the south part of
Scandinavia, it is affected by the overgrowing of higher vegetation.
Conservation
Three localities were taken under monitoring in Estonia, where either grazing or simulation of grazing is needed (by
breaking the sod and opening the soil artificially) in order to re-create and preserve suitable open soil patches for this
species (Vellak and Ingerpuu 2012). Similar management can be recommended elsewhere. Research into threats is also
recommended. It is listed as Endangered in Finland, Vulnerable in Norway and Near Threatened in Sweden (Hodgetts
2015). It is known to occur in protected areas.
Bibliography
Hallingbäck, T., Lönnell, N. and Weibull, H. 2006. Encyclopedia of the Swedish Flora and Fauna: Bladmossor: Sköldmossor–
Blåmossor. Bryophyta: Buxbaumia–Leucobryum. ArtDatabanken, Uppsala.
Hodgetts, N.G. 2015. Checklist and country status of European bryophytes – towards a new Red List for Europe. Irish
Wildlife Manuals, No. 84. National Parks and Wildlife Service, Department of the Arts, Heritage and the Gaeltacht, Ireland.
IUCN. 2019. The IUCN Red List of Threatened Species. Version 2019-2. Available at: www.iucnredlist.org. (Accessed: 04 July
2019).
Kučera, J. and Váňa, J. 2003. Check- and Red List of bryophytes of the Czech Republic. Preslia 75: 193-222.
Vellak, K. and Ingerpuu, N. 2012. The state of bryophyte conservation in Estonia. Studia Botanica Hungarica 43: 59-68.
43
5
44
Appendix 3. Red List status of
European mosses, liverworts and
hornworts
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
ACROBOLBACEAE
Acrobolbus azoricus EN B2ab(iii,iv,v) EN B2ab(iii,iv,v) Yes Yes
Acrobolbus madeirensis EN B1ab(ii,iii,iv,v)+2ab(ii,iii,
iv,v); C2a(i) EN B2ab(ii,iii,iv,v);
C2a(i) Yes Ye s
Acrobolbus wilsonii VU D1 VU D1 Yes No
ADELANTHACEAE
Adelanthus lindenbergianus EN C2a(i) EN C2a(i) No No
Pseudomarsupidium decipiens LC LC No No
AMBLYSTEGIACEAE
Amblystegium serpens LC LC No No
Anacamptodon splachnoides NT B2b(iii) NT B2b(iii) No No
Arvernella microclada EN D EN D Yes Yes
Campyliadelphus chrysophyllus LC LC No No
Campyliadelphus elodes NT A2c NT A2c No No
Campylium laxifolium LC LC No No
Campylium longicuspis VU D1 CR D No No
Campylium protensum LC LC No No
Campylium stellatum LC LC No No
Campylophyllopsis calcarea LC LC No No
Campylophyllopsis sommerfeltii LC LC No No
Conardia compacta NT D1 NT D1 No No
Cratoneuron curvicaule LC LC No No
Cratoneuron licinum LC LC No No
Drepanocladus aduncus LC LC No No
Drepanocladus angustifolius VU B2ab(iii) VU B2ab(iii); C2a (i) No No
Drepanocladus arcticus NT B2b(iii) NE No No
Drepanocladus brevifolius LC NE No No
Drepanocladus capillifolius NT B1b(iii,v) NT No No
Drepanocladus lycopodioides VU A2c; B2ab(ii,iii,iv,v) VU A2c; B2ab(ii,iii,iv,v) No No
Drepanocladus polygamus LC LC No No
Drepanocladus sendtneri VU A2c; B2ab(ii,iii,iv,v);
C2a(i) VU A2c; B2ab(ii,iii,iv,v);
C2a(i) No No
Drepanocladus sordidus NT B2ab(i) NT No No
Drepanocladus trifarius LC LC No No
Drepanocladus turgescens LC VU C2a(i) No No
Hygroamblystegium uviatile LC LC No No
Hygroamblystegium humile LC LC No No
Hygroamblystegium tenax LC LC No No
Hygroamblystegium varium LC LC No No
Hygrohypnella ochracea LC LC No No
Hygrohypnella polaris LC NT No No
45
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Hygrohypnum luridum LC LC No No
Hygrohypnum styriacum EN B2ab(iii); C2a(i) EN B2ab(iii); C2a(i) No No
Leptodictyum riparium LC LC No No
Ochyraea tatrensis CR B1ab(iii,v)+2ab(iii,v);
C2a(i); D CR B1ab(iii)+2ab(iii);
C2a(i); D No No
Palustriella commutata LC LC No No
Palustriella decipiens LC LC No No
Palustriella falcata LC LC No No
Platydictya jungermannioides LC LC No No
Platyhypnum alpestre LC LC No No
Platyhypnum alpinum LC LC No No
Platyhypnum cochlearifolium EN C2a(i) EN C2a (i) No No
Platyhypnum duriusculum LC LC No No
Platyhypnum molle VU C2a(i) VU C2a(i) No No
Platyhypnum norvegicum VU D1 EN D No No
Platyhypnum smithii LC LC No No
Pseudoamblystegium subtile LC LC No No
Pseudocampylium radicale LC LC No No
Pseudohygrohypnum eugyrium LC LC No No
Pseudohygrohypnum subeugyrium NT D1 VU D1 No No
Sanionia nivalis NT B2b(iii) NT No No
Sanionia orthothecioides LC LC No No
Sanionia uncinata LC LC No No
Serpoleskea confervoides LC LC No No
ANASTROPHYLLACEAE
Biantheridion undulifolium EN B2ab(i,ii,iii,iv,v); C2a(i) EN B2ab(i,ii,iii,iv,v);
C2a(i) No No
Crossocalyx hellerianus LC LC No No
Neoorthocaulis attenuatus LC LC No No
Neoorthocaulis binsteadii LC LC No No
Neoorthocaulis oerkei LC LC No No
Neoorthocaulis hyperboreus VU D1 NE No No
Orthocaulis atlanticus LC LC No No
Orthocaulis cavifolius DD DD No No
Schljakovia kunzeana LC LC No No
Schljakovianthus quadrilobus LC LC No No
Tetralophozia liformis CR D CR D No No
Tetralophozia setiformis LC NT No No
ANDREAEACEAE
Andreaea alpestris DD DD Yes No
Andreaea alpina LC LC No No
Andreaea blyttii NT A3c; B2b(iii,iv,v) NT No No
Andreaea crassinervia EN C2a(i) EN C2a(i) No No
Andreaea exuosa EN B1ab(ii,iii,iv,v)+2ab(ii,
iii,iv,v) EN B1ab(ii,iii,iv,v)+2ab(
ii,iii,iv,v) No No
Andreaea frigida VU C1 VU C1 Yes No
Andreaea heinemannii NT B2a NT B2a No No
Andreaea hookeri LC LC No No
46
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Andreaea megistospora LC LC No No
Andreaea mutabilis LC LC No No
Andreaea nivalis NT LC No No
Andreaea rothii LC LC No No
Andreaea rupestris LC LC No No
Andreaea sinuosa VU D1+2 VU D1+2 No No
ANEURACEAE
Aneura latissima EN B2ab(ii,iii) EN B2ab(ii,iii) No No
Aneura maxima DD DD No No
Aneura mirabilis NT D1 NT No No
Aneura pinguis LC LC No No
Riccardia chamedryfolia LC LC No No
Riccardia incurvata LC LC No No
Riccardia latifrons LC LC No No
Riccardia multida LC LC No No
Riccardia palmata LC LC No No
ANOMODONTACEAE
Anomodon attenuatus LC LC No No
Anomodon longifolius LC LC No No
Anomodon rugelii NT D1 VU D1 No No
Anomodon tristis VU D1 VU No No
Anomodon viticulosus LC LC No No
ANTHELIACEAE
Anthelia julacea LC LC No No
Anthelia juratzkana LC LC No No
ANTHOCEROTACEAE
Anthoceros agrestis NT A2c+3c NT A2c+3c No No
Anthoceros caucasicus LC LC No No
Anthoceros neesii EN B2ab(iii) EN B2ab(iii) Yes Yes
Anthoceros punctatus LC LC No No
ARCHIDIACEAE
Archidium alternifolium LC LC No No
ARNELLIACEAE
Arnellia fennica LC NT No No
Gongylanthus ericetorum LC LC No No
Southbya nigrella LC LC No No
Southbya tophacea LC LC No No
AULACOMNIACEAE
Aulacomnium androgynum LC LC No No
Aulacomnium palustre LC LC No No
Aulacomnium turgidum LC LC No No
AYTONIACEAE
Asterella africana VU C2a(i) VU C2a(i) No No
Asterella lindenbergiana LC LC No No
Asterella saccata EN B2ab(iii,iv,v) EN B2ab(iii,iv,v) No No
Mannia androgyna LC LC No No
Mannia californica EN D EN D No No
47
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Mannia controversa EN D EN D No No
Mannia fragrans VU A2c VU A2c No No
Mannia gracilis LC LC No No
Mannia pilosa LC LC No No
Mannia sibirica CR D CR D No No
Mannia triandra VU D1 VU D1 No No
Plagiochasma appendiculatum VU D2 VU D2 No No
Plagiochasma rupestre LC LC No No
Reboulia hemisphaerica LC LC No No
BARTRAMIACEAE
Anacolia menziesii VU D1 VU D1 No No
Anacolia webbii LC LC No No
Bartramia aprica LC LC No No
Bartramia breviseta VU D1 NE No No
Bartramia halleriana LC LC No No
Bartramia ithyphylla LC LC No No
Bartramia laevisphaera EN D EN D No No
Bartramia pomiformis LC LC No No
Bartramia subulata EN D EN D No No
Breutelia azorica EN A3c; B2ab(iii,v) EN A3c; B2ab(iii,v) Yes Yes
Breutelia chrysocoma LC LC Ye s No
Conostomum tetragonum LC LC No No
Philonotis caespitosa LC LC No No
Philonotis calcarea NT A3c NT A3c No No
Philonotis capillaris LC LC No No
Philonotis cernua CR B1ab(i,ii,iii,iv,v)+2ab(i,ii,
iii,iv,v); C2a(i); D CR B2ab(i,ii,iii,iv,v);
C2a(i); D No No
Philonotis fontana LC LC No No
Philonotis hastata NT B2ab(iii) NT B2ab(iii) No No
Philonotis marchica EN C2a(i) EN C2a(i) No No
Philonotis rigida VU C2a(i) VU C2a(i) No No
Philonotis seriata LC LC No No
Philonotis tomentella LC NT No No
Philonotis uncinata VU B2ab(iii) VU B2ab(iii) No No
Plagiopus oederianus LC LC No No
BLASIACEAE
Blasia pusilla LC LC No No
BRACHYTHECIACEAE
Brachytheciastrum collinum LC NT D1 No No
Brachytheciastrum dieckei LC LC No No
Brachytheciastrum olympicum VU B2ab(ii,iv) VU B2ab(ii,iv) No No
Brachytheciastrum trachypodium LC LC No No
Brachytheciastrum vanekii EN B1ab(iii,iv,v)+2ab(iii,iv,v) EN B1ab(iii,iv,v)+
2ab(iii,iv,v) Yes Yes
Brachytheciastrum velutinum LC LC No No
Brachythecium albicans LC LC No No
Brachythecium buchananii EN B2ab(iii); D NE No No
Brachythecium campestre LC LC No No
Brachythecium capillaceum LC DD No No
48
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Brachythecium cirrosum LC LC No No
Brachythecium erythrorrhizon LC LC No No
Brachythecium funkii VU D1 VU D1 Yes Yes
Brachythecium geheebii VU D1 VU D1 Ye s No
Brachythecium glareosum LC LC No No
Brachythecium japygum LC LC Yes No
Brachythecium laetum LC LC No No
Brachythecium mildeanum LC LC No No
Brachythecium novae-angliae LC VU D1 No No
Brachythecium rivulare LC LC No No
Brachythecium rutabulum LC LC No No
Brachythecium salebrosum LC LC No No
Brachythecium tauriscorum LC LC No No
Brachythecium tenuicaule LC LC Yes No
Brachythecium tommasinii LC LC No No
Brachythecium turgidum LC LC No No
Brachythecium udum LC LC No No
Cirriphyllum crassinervium LC LC No No
Cirriphyllum piliferum LC LC No No
Clasmatodon parvulus RE RE No No
Eurhynchiastrum pulchellum LC LC No No
Eurhynchium angustirete LC LC No No
Eurhynchium striatum LC LC No No
Hedenasiastrum percurrens EN A3c EN A3c Yes Yes
Homalothecium aureum LC LC No No
Homalothecium lutescens LC LC No No
Homalothecium mandonii VU A3c VU A3c No No
Homalothecium meridionale LC LC Yes No
Homalothecium philippeanum LC LC No No
Homalothecium sericeum LC LC No No
Kindbergia praelonga LC LC No No
Microeurhynchium pumilum LC LC No No
Myuroclada longiramea DD NA No No
Myuroclada maximowiczii NA NE No No
Nobregaea latinervis EX EX Yes Ye s
Oxyrrhynchium hians LC LC No No
Oxyrrhynchium schleicheri LC LC No No
Oxyrrhynchium speciosum LC LC No No
Palamocladium euchloron EN B2ab(iii); D CR D No No
Platyhypnidium grolleanum DD DD Yes Yes
Pseudorhynchostegiella duriaei NT NT No No
Pseudoscleropodium purum LC LC No No
Rhynchostegiella azorica NT B1a+2a NT B1a+2a Ye s Ye s
Rhynchostegiella bourgaeana EN A3c; B2ab(iii) EN A3c; B2ab(iii) Yes Ye s
Rhynchostegiella curviseta LC LC No No
Rhynchostegiella litorea LC LC No No
49
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Rhynchostegiella pseudolitorea NT B2b(iii) NT B2b(iii) Ye s Yes
Rhynchostegiella tenella LC LC No No
Rhynchostegiella teneriae LC LC No No
Rhynchostegiella trichophylla VU A3c VU A3c Yes Yes
Rhynchostegiella tubulosa DD DD Ye s Ye s
Rhynchostegium alopecuroides LC LC Yes No
Rhynchostegium confertum LC LC No No
Rhynchostegium confusum VU D1 VU D1 Yes Yes
Rhynchostegium megapolitanum LC LC No No
Rhynchostegium murale LC LC No No
Rhynchostegium riparioides LC LC No No
Rhynchostegium rotundifolium LC LC No No
Rhynchostegium strongylense EN D EN D Ye s Yes
Sciuro-hypnum curtum LC LC No No
Sciuro-hypnum dovrense VU C2a(i) EN C2a(i) No No
Sciuro-hypnum otowianum LC LC No No
Sciuro-hypnum glaciale LC LC No No
Sciuro-hypnum latifolium LC NT No No
Sciuro-hypnum oedipodium DD NE No No
Sciuro-hypnum ornellanum EN D EN D No No
Sciuro-hypnum plumosum LC LC No No
Sciuro-hypnum populeum LC LC No No
Sciuro-hypnum reexum LC LC No No
Sciuro-hypnum starkei LC LC No No
Sciuro-hypnum tromsoeense LC LC No No
Scleropodium cespitans LC LC No No
Scleropodium touretii LC LC No No
Scorpiurium circinatum LC LC No No
Scorpiurium deexifolium LC LC No No
Scorpiurium sendtneri LC LC No No
Tomentypnum nitens NT A2c NT No No
BRUCHIACEAE
Bruchia exuosa CR D CR D No No
Bruchia vogesiaca EN B2ab(ii,iii,iv) EN B2ab(ii,iii,iv) No No
Trematodon ambiguus LC LC No No
Trematodon brevicollis VU D1 VU D1 No No
Trematodon laetevirens EN D EN D No No
Trematodon longicollis VU D1 VU D1 No No
Trematodon perssoniorum CR B1ab(iii) CR B1ab(iii) Yes Ye s
BRYACEAE
Anomobryum bavaricum VU D1 VU D1 No No
Anomobryum concinnatum LC LC No No
Anomobryum julaceum LC LC No No
Anomobryum lusitanicum VU D1 VU D1 Yes Yes
Brachymenium notarisii NT B2b(iii,iv,v) NT B2b(iii,iv,v) No No
Brachymenium paradoxum DD NE Yes No
Brachymenium philonotula RE RE No No
50
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Bryum apiculatum LC LC No No
Bryum argenteum LC LC No No
Bryum austriacum VU D1 VU D1 Yes No
Bryum blindii EN D EN D No No
Bryum calophyllum EN B2ab(ii,iii,iv,v) EN B2ab(ii,iii,iv,v) No No
Bryum canariense LC LC No No
Bryum cellulare EN B2ab(iii) EN B2ab(iii) No No
Bryum cryophilum NT B2b(i,ii,iii,iv,v) VU C2a(i) No No
Bryum demaretianum DD DD Yes No
Bryum dichotomum LC LC No No
Bryum dixonii NT D1 NT D1 Yes No
Bryum dyrynense NT B2b(iii,iv,v) NT B2b(iii,iv,v) Yes No
Bryum elegans LC LC No No
Bryum funkii VU B2ab(iii) EN B2ab(iii) No No
Bryum gemmiferum LC LC No No
Bryum gemmilucens LC LC No No
Bryum gemmiparum LC LC No No
Bryum intermedium DD DD No No
Bryum klinggraei LC LC No No
Bryum knowltonii VU C2a(i) VU C2a(i) No No
Bryum kunzei LC LC No No
Bryum marratii EN B2ab(ii,iii,iv,v) EN B2ab(ii,iii,iv,v) No No
Bryum miniatum VU D1 NE No No
Bryum minii LC LC Yes Ye s
Bryum oblongum NT C2a(i) VU C2a(i) No No
Bryum radiculosum LC LC No No
Bryum riparium VU D1 VU D1 No No
Bryum ruderale LC LC No No
Bryum salinum VU C2a(i) EN C2a(i) No No
Bryum sauteri LC LC No No
Bryum schleicheri LC LC No No
Bryum subapiculatum LC LC No No
Bryum tenuisetum LC LC No No
Bryum turbinatum VU C2a(i) VU C2a(i) No No
Bryum valparaisense VU D1 VU D1 No No
Bryum versicolor EN B2b(ii,iii,iv,v)c(iii,iv) EN B2b(ii,iii,iv,v)c(iii,iv) Yes No
Bryum violaceum LC LC No No
Bryum warneum VU B2ab(i,ii,iii,iv,v) VU B2ab(i,ii,iii,iv,v) No No
Bryum weigelii LC LC No No
Bryum wrightii NT VU D1 No No
Imbribryum alpinum LC LC No No
Imbribryum mildeanum LC LC No No
Imbribryum muehlenbeckii LC LC No No
Ptychostomum arcticum LC LC No No
Ptychostomum bornholmense LC LC Yes No
Ptychostomum capillare LC LC No No
51
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Ptychostomum cernuum EN B2ab(i,ii,iii,iv,v) EN B2ab(i,ii,iii,iv,v) No No
Ptychostomum compactum LC LC No No
Ptychostomum creberrimum LC LC No No
Ptychostomum cyclophyllum LC LC No No
Ptychostomum demissum EN C2a(i) EN C2a(i) No No
Ptychostomum donianum LC LC No No
Ptychostomum imbricatulum LC LC No No
Ptychostomum inclinatum LC LC No No
Ptychostomum longisetum CR C2a(i) CR C2a(i) No No
Ptychostomum moravicum LC LC No No
Ptychostomum pallens LC LC No No
Ptychostomum pallescens LC LC No No
Ptychostomum pseudotriquetrum LC LC No No
Ptychostomum rubens LC LC No No
Ptychostomum torquescens LC LC No No
Ptychostomum zieri LC LC No No
Rhodobryum ontariense LC LC No No
Rhodobryum roseum LC LC No No
BRYOXIPHIACEAE
Bryoxiphium madeirense EN A3c EN A3c No No
Bryoxiphium norvegicum LC CR B1ab(iii)+2ab(iii); D No No
BUXBAUMIACEAE
Buxbaumia aphylla LC LC No No
Buxbaumia viridis LC LC No No
CALLIERGONACEAE
Calliergon cordifolium LC LC No No
Calliergon giganteum LC LC No No
Calliergon megalophyllum LC LC No No
Calliergon richardsonii LC LC No No
Hamatocaulis lapponicus EN B2ab(i,ii,iii,iv,v) EN B2ab(i,ii,iii,iv,v) No No
Hamatocaulis vernicosus VU A2c VU A2c No No
Loeskypnum badium LC LC No No
Sarmentypnum exannulatum LC LC No No
Sarmentypnum sarmentosum LC LC No No
Straminergon stramineum LC LC No No
Warnstora uitans LC LC No No
Warnstora procera LC LC No No
Warnstora pseudostraminea LC LC No No
Warnstora trichophylla LC LC No No
Warnstora tundrae LC LC No No
CALYCULARIACEAE
Calycularia laxa CR D NE No No
CALYMPERACEAE
Calymperes erosum CR D CR D No No
CALYPOGEIACEAE
Calypogeia arguta LC LC No No
Calypogeia azorica EN B2ab(ii,iii) EN B2ab(ii,iii) Yes Yes
52
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Calypogeia azurea LC LC No No
Calypogeia ssa LC LC No No
Calypogeia integristipula LC LC No No
Calypogeia muelleriana LC LC No No
Calypogeia neesiana LC LC No No
Calypogeia sphagnicola LC LC No No
Calypogeia suecica LC LC No No
Mnioloma fuscum VU C2a(i) VU C2a(i) No No
CATOSCOPIACEAE
Catoscopium nigritum LC LC No No
CEPHALOZIACEAE
Cephalozia ambigua LC LC No No
Cephalozia bicuspidata LC LC No No
Cephalozia lacinulata CR C2a(i); D CR C2a(i); D No No
Cephalozia macounii CR D CR D No No
Fuscocephaloziopsis anis NT NT No No
Fuscocephaloziopsis albescens LC NT No No
Fuscocephaloziopsis catenulata LC LC No No
Fuscocephaloziopsis connivens LC LC No No
Fuscocephaloziopsis crassifolia LC LC No No
Fuscocephaloziopsis leucantha LC LC No No
Fuscocephaloziopsis loitlesbergeri LC LC No No
Fuscocephaloziopsis lunulifolia LC LC No No
Fuscocephaloziopsis macrostachya LC LC No No
Fuscocephaloziopsis pleniceps LC LC No No
Hygrobiella laxifolia LC LC No No
Nowellia curvifolia LC LC No No
Odontoschisma denudatum LC LC No No
Odontoschisma elongatum LC LC No No
Odontoschisma uitans LC LC No No
Odontoschisma francisci NT C2a(i) VU C2a(i) No No
Odontoschisma macounii LC LC No No
Odontoschisma sphagni LC LC No No
CEPHALOZIELLACEAE
Cephaloziella arctogena VU D1 EN D No No
Cephaloziella aspericaulis CR D DD No No
Cephaloziella baumgartneri LC LC No No
Cephaloziella calyculata NT NT No No
Cephaloziella dentata EN B2ab(iii,v) EN B2ab(iii,v) No No
Cephaloziella divaricata LC LC No No
Cephaloziella elachista VU C2a(i) VU C2a(i) No No
Cephaloziella elegans DD DD No No
Cephaloziella granatensis EN B1ab(iii,iv)+2ab(iii,iv) EN B1ab(iii,iv
+2ab(iii,iv) No No
Cephaloziella grimsulana DD DD No No
Cephaloziella hampeana LC LC No No
Cephaloziella integerrima EN B2ab(iii,v); C2a(i) EN B2ab(iii,v); C2a(i) No No
53
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Cephaloziella massalongi EN B2ab(ii,iii,iv,v) EN B2ab(ii,iii,iv,v) No No
Cephaloziella nicholsonii EN C2a(i) EN C2a(i) Ye s Ye s
Cephaloziella phyllacantha CR D CR D No No
Cephaloziella polystratosa EN D NE No No
Cephaloziella rubella LC LC No No
Cephaloziella spinigera NT B2b(iv,v) NT No No
Cephaloziella stellulifera LC LC No No
Cephaloziella turneri LC LC No No
Cephaloziella uncinata NT B2b(iii,iv) DD No No
Cephaloziella varians LC LC No No
CINCLIDOTACEAE
Cinclidotus aquaticus LC LC No No
Cinclidotus danubicus LC LC No No
Cinclidotus fontinaloides LC LC No No
Cinclidotus riparius LC LC No No
Cinclidotus vivesii DD DD Yes Ye s
CLEVEACEAE
Clevea hyalina LC LC No No
Clevea spathysii NT NT No No
Peltolepis quadrata LC LC No No
Sauteria alpina LC LC No No
CLIMACIACEAE
Climacium dendroides LC LC No No
CONOCEPHALACEAE
Conocephalum conicum LC LC No No
Conocephalum salebrosum LC LC No No
CORSINIACEAE
Corsinia coriandrina LC LC No No
CRYPHAEACEAE
Cryphaea heteromalla LC LC No No
Dendrocryphaea lamyana NT LC Ye s Ye s
CYATHODIACEAE
Cyathodium foetidissimum CR D CR D No No
DALTONIACEAE
Achrophyllum dentatum NA NA No No
Calyptrochaeta apiculata NA NA No No
Daltonia splachnoides LC LC No No
Daltonia stenophylla EN B2ab(iii) EN B2ab(iii) No No
Distichophyllum carinatum CR C2a(i); D CR C2a(i); D No No
DELAVAYELLACEAE
Liochlaena lanceolata LC LC No No
Liochlaena subulata NT NT No No
DICRANACEAE
Aongstroemia longipes LC LC No No
Cnestrum alpestre LC NT No No
Cnestrum glaucescens NT VU D1 No No
Cnestrum schisti LC LC No No
54
Taxonomy IUCN Red List
Category (Europe)
IUCN Red List
Criteria (Europe)
IUCN Red List
Category (EU 28)
IUCN Red List
Criteria (EU 28)
Endemic
to Europe
Endemic
to EU 28
Dicranella cerviculata LC LC No No
Dicranella crispa LC LC No No
Dicranella grevilleana LC LC No No
Dicranella heteromalla LC LC No No
Dicranella howei LC LC No No
Dicranella humilis LC LC No No
Dicranella rufescens LC LC No No
Dicranella schreberiana LC LC No No
Dicranella staphylina LC LC No No
Dicranella subulata LC LC No No
Dicranella varia LC LC No No
Dicranum acutifolium LC LC No No
Dicranum angustum LC LC No No
Dicranum bardunovii DD NE No No
Dicranum bonjeanii LC LC No No
Dicranum brevifolium LC LC No No
Dicranum crassifolium NT B2b(i) LC Ye s Ye s
Dicranum dispersum EN D EN D No No
Dicranum drummondii LC NT No No
Dicranum elongatum LC LC No No
Dicranum agellare LC LC No No
Dicranum exicaule LC LC No No
Dicranum fragilifolium LC NT No No
Dicranum fulvum LC LC No No
Dicranum fuscescens LC LC No No
Dicranum groenlandicum LC NT No No
Dicranum laevidens LC VU D No No
Dicranum leioneuron LC LC No No
Dicranum majus LC LC No No
Dicranum montanum LC LC No No
Dicranum muehlenbeckii VU C2a(i) VU C2a(i) No No
Dicranum polysetum LC LC No No
Dicranum schljakovii DD NE No No
Dicranum scoparium LC LC No No
Dicranum scottianum LC LC Yes No