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Bryophyte conservation on a North Atlantic hotspot:
threatened bryophytes in Madeira and Selvagens
Archipelagos (Portugal)
Manuela Sim-Simab, Sara Ruasab, Susana Fontinhac, Lars Hedenäsd, Cecília Sérgiob & Carlos
Loboe
a Universidade de Lisboa, Faculdade de Ciências de Lisboa, DBV, Centro de Biologia
Ambiental, C2, Campo Grande, 1749-016 Lisboa, Portugal
b Museu Nacional de História Natural e da Ciência, Jardim Botânico/CBA, Rua da Escola
Politécnica, n° 58, 1250-102 Lisboa, Portugal
c ISOPlexis Universidade da Madeira/SRA, Campus da Penteada, 9000-390 Funchal, Madeira,
Portugal
d Swedish Museum of Natural History, Department of Botany, Box 50007 SE-104 05
Stockholm, Sweden
e Jardim Botânico da Madeira – Eng. Rui Vieira, Quinta do Bom Sucesso, 9064-512 Funchal,
Madeira, Portugal
Published online: 27 May 2014.
To cite this article: Manuela Sim-Sim, Sara Ruas, Susana Fontinha, Lars Hedenäs, Cecília Sérgio & Carlos Lobo (2014):
Bryophyte conservation on a North Atlantic hotspot: threatened bryophytes in Madeira and Selvagens Archipelagos (Portugal),
Systematics and Biodiversity, DOI: 10.1080/14772000.2014.918063
To link to this article: http://dx.doi.org/10.1080/14772000.2014.918063
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Research Article
Bryophyte conservation on a North Atlantic hotspot: threatened
bryophytes in Madeira and Selvagens Archipelagos (Portugal)
MANUELA SIM-SIM
1,2
*, SARA RUAS
1,2
, SUSANA FONTINHA
3
, LARS HEDEN
€
AS
4
, CEC
ILIA S
ERGIO
2
& CARLOS LOBO
5
1
Universidade de Lisboa, Faculdade de Ci^
encias de Lisboa, DBV, Centro de Biologia Ambiental, C2, Campo Grande, 1749-016 Lisboa,
Portugal
2
Museu Nacional de Hist
oria Natural e da Ci^
encia, Jardim Bot^
anico/CBA, Rua da Escola Polit
ecnica, n58, 1250-102 Lisboa, Portugal
3
ISOPlexis Universidade da Madeira/SRA, Campus da Penteada, 9000-390 Funchal, Madeira, Portugal
4
Swedish Museum of Natural History, Department of Botany, Box 50007 SE-104 05 Stockholm, Sweden
5
Jardim Bot^
anico da Madeira Eng. Rui Vieira, Quinta do Bom Sucesso, 9064-512 Funchal, Madeira, Portugal
(Received 5 February 2014; revised 17 March 2014; accepted 31 March 2014)
On oceanic islands, the evolution of plants and animals with particular characteristics is favoured due to their isolation,
populations normally comprising a large number of unique, endemic species. The Madeira and Selvagens archipelagos are
considered biodiversity hotspots, containing an especially rich bryoflora. Due to its characteristically small size, this
taxonomic group does not get much attention in conservation programmes. However, these plants are an important
component of terrestrial ecosystems, representing a major part of biodiversity and playing a vital role in the ecosystem’s
functioning. As such, the development of the first Red List for Madeira and Selvagens Archipelagos has the potential to
guide conservation efforts focused on taxa and habitats where threatened species and endemics are better represented. By
applying the International Union for Conservation of Nature and Natural Resources (IUCN) criteria, recently revised to
apply to small islands, it was possible to obtain the percentage of threatened taxa present in the archipelagos (23.6%), and
for each habitat type. It was verified that high mountain habitats and the Laurel forest represent areas that host higher
percentages of threatened taxa (29.5% and 22.2%). An important result of the present Red List is the identification of
hotspots for bryophyte diversity, supporting the definition of reserves/microreserves. The information obtained can also be
linked up with the Red Lists of other taxonomic groups to work towards the definition of a more holistic conservation
strategy.
Key words: biodiversity, bryophytes, conservation, IUCN criteria, Madeira region, Red List
Introduction
Oceanic islands are tiny dots in the middle of the ocean,
but they are of great importance to the understanding of
many evolutionary and ecological processes (e.g. Gilles-
pie et al.,2008; Kueffer & Fern
andez-Palacios, 2010).
Due to their isolation they favour the evolution of plants
and animals with particular characteristics, comprising a
large number of unique species (Witt & Maliakal-Witt,
2007). The distinct altitudinal levels that frequently occur
on volcanic oceanic islands (Yu & Lei, 2001) result in a
very limited capability on the part of species to extend
their ranges, making them more vulnerable to changes in
environmental conditions, compared with continental
areas (Lowry II, 2002). Bryophytes are more easily dis-
persed than many other organisms, thanks to their small
spores, and could therefore potentially disperse to and
from isolated islands relatively easily, and be less affected
by the specific conditions of them. However, recent stud-
ies have shown that oceanic island bryophytes also display
island syndromes to a significant extent (Pati~
no et al.,
2013a), and that their diversity on such islands is predi-
cated mainly by habitat availability and complexity rather
than island age and distance to the closest continent
(Pati~
no et al.,2013b).
The islands of Madeira, Porto Santo, Desertas and Sel-
vagens, along with other Macaronesian archipelagos, are
included in the Mediterranean biodiversity hotspots
(Myers et al.,2000), and Madeira Island stands out as the
second richest island of all the Atlantic archipelagos of
*Correspondence to: Maria Manuela Pinheiro Sim-Sim. E-mail:
msimsim@fc.ul.pt
ISSN 1477-2000 print/ 1478-0933 online
ÓThe Trustees of the Natural History Museum, London 2014. All Rights Reserved.
http://dx.doi.org/10.1080/14772000.2014.918063
Systematics and Biodiversity (2014), 0(0):116
Downloaded by [Susana Fontinha] at 10:07 29 May 2014
the Azores, Madeira, Selvagens, Canary Islands and Cape
Verde, only surpassed by Tenerife in the Canary Islands
(Izquierdo et al.,2001; Gonz
alez-Mancebo et al.,2013),
the latter being an island which, however, is much larger.
The Madeira and Selvagens archipelagos are located in
the Atlantic Ocean, at the point of intersection between
the tropics, North America and Europe. Madeira, in par-
ticular, harbours a high diversity of habitats where bryo-
phytes occur, from sea level to the highest mountains,
playing an important role in the colonization, stability and
dynamics of its ecosystems (Fontinha et al.,2006,2011;
Michel et al.,2012).
Capelo et al. (2007) posit that the climate of Madeira
Islands includes two main types: Mediterranean and tem-
perate.Onthesouthside,below1000mandonthelow-
lands near the north coast, the climate is characterized by a
long dry season during the summer months. Above 1000 m
on the south side and on most of the north side, the climate
is temperate (Prada et al.,2009). The type of vegetation
that occurs in this area is characterized by a diverse under-
storey (moist Laurisilva). At higher altitudes marked levels
of thermal variation take place, with high temperatures in
summer and very low temperatures in winter, which
explains the absence of laurel type forests in those areas
and the presence of high-altitude scrubland predominated
by Erica spp. (Capelo et al.,2007). Different bryophyte
communities are present at these altitudinal levels and,
according to S
ergio et al.(2008), the bryoflora of the
Madeiran and Selvagens Archipelagos consists of 529 taxa,
with the highest diversity (511 taxa) on Madeira Island.
Bryophytes are inherently small and inconspicuous
plants which do not get much attention in conservation
programmes (Tan & P
ocs, 2000). However, this plant
group is an important component of terrestrial ecosys-
tems, representing a major part of biodiversity in humid
forests, mountains and even coastal habitats, and playing
a vital role in the functioning of ecosystems (Vitt &
Wieder, 2009). As such, Red List data constitute an essen-
tial source of information for guiding conservation efforts
focused on species and the habitats where they occur.
In recent years, the International Union for the Conser-
vation of Nature (IUCN) Red List categories and criteria
have been extensively used at national and regional levels
to evaluate threatened bryophytes, due to the publication
of guidelines for national and regional Red Lists
(G€
ardenfors et al.,2001). Furthermore, species distribu-
tion areas are most often politically divided, which is why
local or regional Red Lists are important in the implemen-
tation of protection strategies by national and local author-
ities (Sabovljevi
cet al.,2004).
In their recent paper, Gonz
alez-Mancebo et al. (2012)
discussed the application of IUCN criteria to bryophytes
in small and highly diverse habitats, and developed a Red
List for the bryophytes of the Canary Islands. Ah-Peng
et al.(2012) provided a regional IUCN Red List for the
bryophytes of R
eunion (the Mascarene archipelago), rep-
resenting the first regional Red List of bryophytes for
Africa. These recent publications suggest new methodolo-
gies and guidelines to apply the IUCN criteria to islands
and small regions. In fact, as noted by Gonz
alez-Mancebo
et al. (2012), the application of the IUCN criteria to bryo-
phytes on islands is associated with some problems due to
different considerations for fragmentation, extent of
occurrence and area of occupation, the distance between
locations and the high diversity of island habitats. Never-
theless, during the past 20 years the advances in research
on the bryoflora of Madeira have facilitated a greater
knowledge of bryophyte distribution in relation to habitat
quality and characteristics, allowing for the improved
application of IUCN criteria to the Madeira and Selvagens
Archipelagos.
Until now, only the rarest, often riparian, taxa and bryo-
phytes with biogeographic singularities from Madeira had
been evaluated (S
ergio et al.,1992;Lu
ıs et al.,2008;
Sim-Sim et al.,2008). The present paper aims to provide
the first Red List of bryophytes for the Madeira region,
identifying threatened taxa and focusing on the habitats
with the highest number of threatened bryophytes. It is
also our aim to indicate areas which are important for
bryophyte diversity and conservation.
Materials and methods
Study area
The volcanic Archipelagos of Madeira (Madeira, Porto
Santo, Desertas) and Selvagens are located in the Atlantic
Ocean. Madeira is the largest island of the Archipelago
(c. 737 km
2
) followed by Porto Santo (42 km
2
), with
Deserta Grande the smallest (c.10km
2
). The Archipelago
of Selvagens constitutes the southernmost part of the Por-
tuguese territory and is situated approximately 300 km
south of Madeira.
The climate in the Madeira Archipelago is considered
to be temperate oceanic, with a well-balanced year-round
temperature, and is mainly regulated by a branch of the
Gulf Stream and by a high-pressure zone called the
‘Azores anticyclone’ (K€
urschner et al., 2007a; Prada
et al.,2009). Clouds, mist and fog are common phenom-
ena, especially in the highest and central parts of Madeira
Island (Fontinha et al.,2006, 2010; Prada et al.,2009).
The laurel forests, on slopes exposed to the prevailing NE
winds, receive more than 3000 mm of rainfall per year,
and have high levels of relative humidity (7590%)
nearly all year round (S
ergio et al.,2008; Prada et al.,
2009; Fontinha et al.,2010). The highly variable topogra-
phy that characterizes Madeira Island in particular has an
important influence on the climate and on the distribution
of vegetation.
2 M. Sim-Sim et al.
Downloaded by [Susana Fontinha] at 10:07 29 May 2014
The values for ‘average annual precipitation’ and
‘average annual temperature’ are sourced from PRAM
(2002) and are the following: Madeira Island 640 mm
and 19 C; Porto Santo 390 mm and 19 C; Desertas
500 mm and 18.5 C; Selvagens 30.6 mm and 19.5 C.
Data on bryophyte presence and distribution
As Hern
andez & Navarro (2007) stated, the data available
in herbarium collections can prove essential for the defini-
tion of threat categories and enable a more accurate use of
IUCN criteria when a Red List is being developed. Signif-
icant numbers of Madeiran bryophytes collected more
than 50 years ago exist, with few of their specimens col-
lected recently. As such, an important starting point for
the development of this Red List was to perform a revi-
sion of the specimens present in the herbaria that contain
most of the Madeira bryophyte collections (LISU,
MADS, MADJ and S). Based on this revision it was possi-
ble to confirm the occurrence of specific taxa and to
exclude from Madeira bryophytes a few that had been
included owing to erroneous identifications or old taxo-
nomic criteria.
In addition to herbarium data, a significant part of the
required information about bryophyte distribution, ecol-
ogy, habitat requirements and life strategies was obtained
from the extensive studies that have been performed in all
Madeira and Selvagens Archipelagos in the past 20 years
(e.g. Heden€
as, 1992; Fontinha & S
ergio, 1993; ECCB,
1995;K€
urschner et al., 2007b,2008a;Lu
ıs et al.,2008,
2012; Sim-Sim et al.,2011). Furthermore, an extensive
bibliographic revision was carried out to complete our
knowledge of distributions and descriptions (e.g. Luisier,
19171922, 1927, 19301932; Persson, 1939;N
obrega,
1990;S
ergio et al.,2006). In some cases it was not possi-
ble to confirm bryophyte occurrence or provide accurate
figures for their frequency and, in such cases, we included
the taxon in the Data Deficient (DD) category.
The nomenclature followed in this assessment is mod-
elled on that used in Ros et al. (2007) for liverworts and
hornworts and Ros et al. (2013) for mosses.
Categories considered
The categories and subcategories considered in this Red
List are based on those of IUCN (2001,2003), Hodgetts
(2000), S
ergio et al. (2007,2013) and the IUCN Standards
and Petitions Subcommittee (2011). Threat categories
include Critically Endangered (CR), Endangered (EN)
and Vulnerable (VU). The category Near Threatened
(NT) is considered when a taxon does not yet qualify as
threatened, but may be close to qualifying as threatened.
In the present Red List it was also applied in the following
scenarios: when the maintenance of the taxon status is
dependent on the conservation and integrity of the habitat
(IUCN, 2001; Schnyder et al.,2004; IUCN Standards and
Petitions Subcommittee, 2011) when the taxon has a very
restricted distribution; when there are no current threats,
but there are plausible events that may cause decline
(IUCN Standards and Petitions Subcommittee, 2011); and
when the taxon is an endemic to Macaronesia or Madeira.
A taxon is considered to be Extinct (EX) when there is
no reasonable doubt that the last individual has disap-
peared and is Regionally Extinct (RE) in Madeira, if there
are no records after 1960 and all known localities have
been visited and surveyed without success. The Data Defi-
cient (DD) category is considered for: recently discovered
taxa (DD-n subcategory); taxa which can only be identi-
fied by experts or by using complex techniques; taxa that
have occurred possibly as ‘chance’ ephemerals; taxa with
inadequate information to make an assessment of their
risk of extinction based on their distribution and/or popu-
lation status; several taxa that have not been seen for over
50 years but which cannot be regarded as extinct because
historical localities were insufficiently investigated (DD-
va subcategory) (Hodgetts, 2000; IUCN, 2001,2003;
IUCN Standards and Petitions Subcommittee, 2011;
S
ergio et al.,2007,2013).
Criteria applied
The criteria applied follow the guidelines (IUCN, 2001;
IUCN, 2003; IUCN Standards and Petitions Subcommit-
tee, 2011) and include the relevant adaptations of criterion
B to small islands by Gonz
alez-Mancebo et al. (2012)
namely: (i) AOO (Area of Occupancy) was reduced to
0.25 km
2
(grid of 0.5 km 0.5 km) to avoid overestima-
tion of the size of suitable habitats; (ii) New AOO thresh-
olds were estimated (taking into consideration this
smaller scale) by analysing the frequency for all bryo-
phytes present in Madeira (excluding DD and doubtful
occurrences). Using Gonz
alez-Mancebo et al. (2012)
methodology, a 50th percentile of 24 grids was obtained
(corresponding to the Near Threatened category), and
from a reduction of one-quarter for each following threat
category, resulting in six grid squares for the Vulnerable
category, two grid squares for Endangered and one grid
square for Critically Endangered; (iii) Location size: the
location size is also expressed in terms of the number of
grid squares for criterion B2 and D2.
To qualify for any of the threat categories on the Red
List, a taxon is required to fulfil a minimum of one crite-
rion (A to E) (IUCN, 2001). Every bryophyte evaluated
was tested against all criteria and was allocated to the high-
est category (IUCN, 2001; IUCN Standards and Petitions
Subcommittee, 2011). The criteria applied were essentially
the B2 criterion and D2, except for Echinodium setigerum
(Mitt.) Jur. and Radula jonesii Bouman et al. (criterion D1
and C2a(i) respectively) (Sim-Sim, 2008).
Threatened bryophytes in Madeira and Selvagens Archipelagos 3
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Criterion B: To qualify for criterion B, the general
distributional threshold must first be met, either in terms
of extent of occurrence (EOO) (not applied) or area of
occupancy (AOO) (e.g. IUCN, 2001; IUCN Standards
and Petitions Subcommittee, 2011). For the Madeiran
bryoflora we considered that AOO is generally better cor-
related with population size than EOO, providing more
accurate estimates of extinction probability based on
changes in within-range habitat extent, fragmentation or
suitability (Joseph & Possingham, 2008).
The taxon must then meet at least two of the three
options listed for criterion B: (a) be severely fragmented
or known to exist in no more than a specific number of
locations; (b) be characterized by continuing decline
(observed, inferred or projected) in at least one of the fol-
lowing: (i) extent of occurrence, (ii) area of occupancy,
(iii) habitat area and or quality and/or (iv) number of loca-
tions or subpopulations; or (c) be characterized by
extreme fluctuation (not applied in this study) (IUCN
Standards and Petitions Subcommittee, 2011). When the
causes of a taxon’s decline were identified we took into
consideration the IUCN Threats Classification Scheme
(Version 3.1) (IUCN, 2012). The evaluation of projected
decline was carried out drawing on published reports and
models on climate change impacts for Madeira ecosys-
tems (see Cruz et al.,2009), land use alterations as well as
the authors’ field experience.
The assessment of potential severe fragmentation for
bryophytes is not always simple. As Hallingb€
ack et al.
(1996,1998) indicate, bryophyte distribution is naturally
patchy as they are frequently confined to specific micro-
sites. Considering that Madeira Island is only 58 km long,
it was fundamental to take into account the island’s highly
irregular geomorphology (natural barriers to dispersal).
According to Pharo & Zartman (2007) taxa that invest
more in competitive ability (e.g. long-lived, large perenni-
als with larger spores), despite being abundant in
undisturbed places, run greater risks of extinction in frag-
mented landscapes compared with more mobile taxa. It
was as such that, for bryophytes whose populations were
markedly separated by a geomorphological barrier and
characterized by weak dispersal mechanisms (dispersal
limited at local scales), fragmentation was considered.
This assessment had to be carried out on case-by-case
bases and taking into account the possibility of gene flow
between patches (Hallingb€
ack et al.,1996).
Criterion D: Criterion D identifies very small or
restricted populations (IUCN, 2001), including subcriterion
D1 and D2. The subcriterion D1 (number of mature indi-
viduals) is not commonly used for bryophytes, given that
in most cases the number of individuals cannot be esti-
mated with sufficient accuracy (Hodgetts, 2000). The D2
subcriterion has been used extensively to assign bryophytes
to VU (5 or fewer locations) (Hodgetts, 2011) and takes
into consideration the fact that the population is prone to
the effects of human activities or stochastic events over a
very short time period with an uncertain future, and is thus
capable of becoming Critically Endangered or even Extinct
in a very short time span (IUCN, 2001).
When applying both criteria (summarized in Table 1)to
evaluate taxa category, besides the quantitative approach,
special attention was also given to taxon-specific informa-
tion, requirements and habitat vulnerability.
Vegetation and habitat type definitions
Capelo et al. (2004) identified six vegetation series for
Madeira Island with which we were able to associate bryo-
phyte distributions, resulting in a simplified habitat designa-
tion based on the recent publication of Gonz
alez-Mancebo
et al. (2013)(Table 2). We considered bryophyte occurrence
in two additional habitat types, namely: anthropogenic/dis-
turbed habitats (artificial walls, roads and trails, etc.); and
freshwater habitats (stream margins, waterfalls, etc.).
Table 1. Categories and criteria used in the present bryophyte Red List of the Madeira and Selvagens archipelagos, except for
Echinodium setigerum (D1) and Radula jonesii (C2ai) (see text of section 2.4).
Threat category
Criteria description CR EN VU NT
B2 Area of Occupancy (AOO) small and
estimations indicating at least two of ab:
(a) Severely fragmented or no more than the
estimated number of locations
(b) Continuing decline
i) Extent of occurrence
ii) Area of occupancy
iii) Quality of habitat
iv) Nof locations or subpopulations
1grid squares 2grid squares 36grid squares 24 grid squares
D2 Population with a very restricted area of
occupancy or number of locations; Taking into
account Nno. of localities or AOO
5grid squares 8grid squares
4 M. Sim-Sim et al.
Downloaded by [Susana Fontinha] at 10:07 29 May 2014
Coastal areas in Madeira, characterized by an arid to
semi-arid harsh environment host a highly typical bryo-
phyte community with specific life strategies (K€
urschner
et al.,2008a; Fontinha et al.,2011). On Madeira Island
this type of environment is mainly restricted to the south-
ern side of the island and occupies a relatively small area.
Lower altitude vegetation, in particular the Olea series-
associated types, is very rare nowadays, and only a very
small percentage is covered by legal protection. Porto
Santo, Desertas and Selvagens are primarily characterized
by dry, exposed environments with an important oceanic
influence (K€
urschner et al.,2008b). Desertas and Selva-
gens are small, uninhabited islands, so human activity
does not directly affect bryophyte population integrity.
However, precisely due to their small size, stochastic
events represent an important threat. In fact, some bryo-
phyte populations are so small that they can easily be
wiped out by a single event.
Mesic areas in the lower portions of the main island are
mainly characterized by the presence of disturbed Semele
androgynaeApollonio barbujanae formations and tran-
sition patches to the Clethro arboreaeOcoteo foetentis
sigmetum (Capelo et al.,2004).
The natural vegetation of this intermediate altitudinal
belt (between the Laurel forest sensu stricto and coastal
areas) is currently highly fragmented, since most of the
agriculture fields, exotic tree plantations and pastures are
located in this area.
Madeira’s area of Laurel forest is currently the best pre-
served of all the Macaronesian islands, covering about
20% of the main island territory (S
ergio et al.,2008). It is
characterized by a constant high level of air humidity and
provides stable shade/moisture conditions, particularly
important for a large proportion of the bryophytes that are
present in this ecosystem. Bryophytes are distributed
across varying micro-habitats and form a significant com-
ponent of the biota, sometimes with close to 100% cover
on tree bark or moist slopes (Fontinha et al.,2006;
K€
urschner et al.,2007a).
Ericaceous forest and thickets are present on steep, sunny
and nutrient-poor exposed slopes, and are characterized by
more marked climatic variations and higher continentality
in comparison with the Clethro arboreaeOcoteetum foe-
tentis (Laurel forest s.l.) (Guimar~
aes & Olmeda, 2008). This
formation occurs particularly frequently at heights above
1400 m where dense E. arborea stands become clearly dom-
inant (Capelo et al.,2004;K
€
urschner et al.,2007a).
High mountain habitats in Madeira Archipelago are
restricted to a small area of the main island, occurring above
1650 m, where there is a predominance of rupicolous herba-
ceous vegetation and small shrub communities and an
almost complete absence of trees (Capelo et al.,2004). This
is probably one of the most threatened habitats in Madeira,
essentially due to landslides, invasive species and to
Madeiran climate change scenarios (Cruz et al.,2009).
Freshwater habitats are mainly represented by cascades,
Levadas (water channels) and slopes along the Levadas, ver-
tical walls with constant dripping water, torrential streams,
stream margins and temporary ponds (Lu
ıs et al.,2010). It
is as such that species which occur along the margins of a
freshwater body, or that are closely associated to water pres-
ence but only temporarily immersed are also considered,
and not only exclusively aquatic species. With urban devel-
opment, increased levels of tourism and agricultural activi-
ties, there has been an increase in water demand, resulting
in lower flow rates in natural water courses, as well as a
decline in water quality (PRAM, 2002), which is aggravated
by climate change consequences (Cruz et al.,2009).
The SPSS programme version 16.0. was used to per-
form a Chi-squared test (x
2
) to understand the variation of
threatened taxa frequency between the areas of Natura
2000 and the areas not protected by legislation.
We hereby certify that, during the field work, all bio-
logical samples collected and transported by the authors
were previously and officially authorized by the Director
of the Natural Park Services (Secretaria Regional do
Ambiente e Recursos Naturais Regi~
ao Aut
onoma da
Madeira, Governo Regional da Madeira).
Table 2. Vegetation and habitat type definition for bryophyte community analysis (Capelo et al.,2004).
Phytosociological nomenclature Habitat type
Mayteno umbellataeOleo maderensis sigmetum (0200 m South slope)
Helichryso melaleuciSideroxylo marmulanae sigmetum
(200300 m South slope); (080 m North slope)
Coastal/lowland habitats
Semele androgynaeApollonio barbujanae and Clethro arboreaeOcoteo
foetentis sigmetum disturbed areas. (300600 m m South slope);
(50450 m North slope)
Mesic areas in lower parts of the Island
Clethro arboreaeOcoteo foetentis sigmetum (8001450 m South slope);
(3001400 North slope)
Laurel forest
Polysticho falcinelliErico arboreae sigmetum (from 1400 to 1650 m) Ericaceous forest and thickets of Erica arborea L.
Armerio maderensisParafestuco albidae microgeosigmetum
(from 1650 m)
High mountain habitat
Threatened bryophytes in Madeira and Selvagens Archipelagos 5
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Results
Composition of the moss flora
The bryoflora of Madeira, based on current taxonomic
concepts and levels of knowledge, contains 538 taxa, 6
hornworts, 172 liverworts and 360 mosses. Endemic taxa
(considering Madeiran endemics and Macaronesian
endemics occurring in the Madeira and Selvagens Archi-
pelagos) represent about 7% of all taxa.
According to our surveys, and to our current knowl-
edge, we consider two Madeiran endemics Extinct (EX)
(Fissidens microstictus Dixon & Luisier and Nobregaea
latinervis Heden€
as). In visits to the known localities of
these species we were not able to locate the plants. The
habitat of F. microstictus is totally urbanized and that of
N. latinervis lies on the edge of the Laurel forest. Two
species are now Regionally Extinct (RE) (Brachymenium
philonotula Broth. and Fissidens fontanus (Bach. Pyl.)
Steud.).
An effort has been made to distinguish between taxa
that have probably genuinely disappeared and those that
may still exist, the latter being recorded under the Data
Deficient category. Excluded taxa and notes on bryo-
phytes with doubtful taxonomic status are illustrated after
the list of threatened taxa (Table 3). In this paper we only
published the threatened and the Near Threatened
Table 3. List of threatened (CR, EN, VU) and Near Threatened
taxa.
Critically Endangered
Taxa Criteria
Fissidens ovatifolius B2ab(ii)(iii)
Gymnocolea inflata B2ab(iii)
Oxyrrhynchium schleicheri B2ab(ii)(iii)
Palustriella commutata B2ab(ii)(iii)
Schistidium strictum B2ab(iii)
Endangered
Taxa Criteria
Aphanolejeunea sintenisii B2ab(iii)
Bryoerythrophyllum inaequalifolium B2ab(iii)
Bryoxiphium madeirense
b
B2ab(ii)(iii)
Calypogeia sphagnicola B2ab(iii)
Cephalozia catenulata B2ab(ii)(iii)
Colura calyptrifolia B2ab(ii)(iii)
Dicranella rufescens B2ab(ii)(iii)
Exsertotheca crispa B2ab(ii)(iii)
Fissidens polyphyllus B2ab(ii)(iii)
Hymostylium recurvirostrum B2ab(iii)
Marsupella profunda
c
B2ab(iii)
Odontoschisma denudatum B2ab(ii)(iii)
Orthodontium gracile B2ab(iii)
Pleurozium schreberi B2ab(ii)(iii)
Pohlia nutans B2ab(iii)
Radula jonesii
b
C2a(i)
Scapania subalpina B2abiii)
Table 3. (Continued )
Sciuro-hypnum populeum B2abiii)
Sphagnum compactum B2ab(ii)(iii)
Syntrichia fragilis B2ab(iii)
Syntrichia princeps B2ab(iii)
Weissia longifolia B2ab(iii)
Vulnerable
Taxa Criteria
Acaulon muticum D2
Acaulon triquetrum D2
Acrobolbus wilsonii B2ab(iii)
Adelanthus decipiens B2ab(ii)(iii)
Alophosia azorica
a
B2ab(iii)
Amphidium curvipes
a
B2ab(iii)
Andreaea flexuosa subsp. luisieri B2ab(ii)(iii)
Andreaea heinemannii subsp. heinemannii D2
Andreaea rupestris B2ab(ii)(iii)
Antitrichia californica D2
Aphanolejeunea azorica
a
B2ab(ii)(iii)
Aphanolejeunea madeirensis
a
B2ab(iii)
Brachymenium notarisii
a
B2ab(iii)
Bryoerythrophyllum campylocarpum
c
B2ab(ii)(iii)
Calypogeia muelleriana B2ab(ii)
Campylostelium strictum B2ab(ii)(iii)
Cephalozia crassifolia B2ab(iii)
Cephalozia lunulifolia B2ab(iii)
Vulnerable
Cephaloziella baumgartneri B2ab(iii)
Cephaloziella dentata B2ab(ii)(iii)
Cephaloziella granatensis B2ab(iii)
Cephaloziella hampeana B2ab(iii)
Cephaloziella rubella B2ab(iii)
Chiloscyphus pallescens B2ab(ii)(iii)
Cololejeunea calcarea D2
Cololejeunea schaeferi
a
B2ab(ii)(iii)
Crossidium crassinerve D2
Ditrichum subulatum B2ab(iii)
Echinodium setigerum
b
D1
Echinodium spinosum
a
B2ab(iii)
Encalypta vulgaris D2
Entosthodon commutatus B2ab(iii)
Ephemerum serratum B2ab(iii)
Eurhynchium striatum B2ab(ii)(iii)
Fissidens nobreganus
b
B2ab(iii)
Fontinalis antipyretica B2abii)iii)
Frullania sergiae
b
D2
Grimmia donniana B2ab(ii)(iii); D2
Grimmia funalis B2ab(ii)(iii); D2
Grimmia montana B2ab(ii)(iii); D2
Grimmia ovalis B2ab(ii)(iii); D2
Gymnostomum aeruginosum B2ab(ii)(iii)
Hedenasiastrum percurrens
b
B2ab(iii)
Hedwigia stellata B2ab(iii)
Homalothecium aureum B2ab(ii)
(continued)
6 M. Sim-Sim et al.
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Table 3. (Continued )
Hygrobiella laxifolia B2ab(ii)
Hyocomium armoricum D2
Hypnum cupressiforme var. lacunosum B2ab(ii)(iii)
Imbribryum mildeanum D2
Isothecium myosuroides B2ab(ii)(iii)
Jungermannia atrovirens B2ab(ii)(iii)
Jungermannia leiantha B2ab(iii)
Leiocolea bantriensis D2
Lejeunea canariensis
a
B2ab(iii)
Lejeunea mandonii B2ab(ii)(iii)
Leptobryum pyriforme B2ab(iii)
Leptodon longisetus
b
B2ab(iii)
Leptoscyphus cuneifolius B2ab(iii)
Leucodon canariensis
a
B2ab(iii)
Lophocolea minor B2ab(iii)
Lophozia bicrenata B2ab(iii)
Lophozia sudetica B2ab(ii)(iii)
Marchantia paleacea B2ab(ii)(iii)
Marsupella funckii B2ab(iii)
Marsupella sprucei B2ab(iii)
Metzgeria conjugata B2ab(iii)
Microcampylopus laevigatus B2ab(ii)(iii)
Nardia geoscyphus B2ab(ii)(iii)
Nowellia curvifolia B2ab(iii)
Vulnerable
Orthodontium pellucens B2ab(iii)
Palustriella falcata D2
Philonotis hastata B2ab(iii)
Plagiochila spinulosa B2ab(iii)
Plagiomnium medium B2ab(ii)(iii)
Pogonatum urnigerum B2ab(iii)
Pohlia annotina B2ab(iii)
Porella cordaeana B2ab(ii)(iii); D2
Porella inaequalis B2ab(iii)
Pseudorhynchostegiella duriaei B2ab(iii)
Racomitrium affine B2ab(iii)
Rhytidiadelphus loreus B2ab(iii)
Rhytidiadelphus triquetrus B2ab(iii)
Riccia atlantica
b
B2ab(iii)
Riccia ligula D2
Scapania curta B2ab(iii)
Scapania umbrosa B2ab(iii)
Scorpiurium deflexifolium B2ab(iii)
Southbya nigrella D2
Sphagnum auriculatum D2
Sphagnum subnitens D2
Syntrichia bogotensis B2ab(iii)
Thamnobryum fernandesii
b,c
B2ab(ii)(iii)
Tortella limbata
a
B2ab(ii)
Tortula canescens B2ab(ii)
Tortula cuneifolia B2ab(ii)(iii)
Tortula lindbergii B2ab(ii)(iii)
Tortula subulata B2ab(iii)
Table 3. (Continued )
Tortula truncate B2ab(ii)(iii)
Tylimanthus madeirensis
b
B2ab(iii)
Zygodon forsteri B2ab(ii)
Near Threatened
Taxa
Aloina aloides
Aloina ambigua
Aloina rigida
Andoa berthelotiana
a
Andreaea alpestris
Aneura pinguis
Anthoceros agrestis
Atrichum androgynum
Atrichum angustatum
Bazzania trilobata
Brachytheciastrum velutinum
Brachythecium albicans
Brachythecium rutabulum var. atlanticum
b
Cinclidotus fontinaloides
Dichodontium pellucidum
Dicranum fuscescens
Dicranum scottianum
Ditrichum flexicaule
Near Threatened
Ditrichum punctulatum
Entosthodon pulchellus
Exsertotheca intermedia
a
Fabronia pusilla
Fissidens coacervatus
Fissidens crassipes subsp. warnstorfii
Fissidens sublineaefolius
b
Frullania polysticta
Grimmia decipiens
Grimmia torquata
Hedwigia ciliata
Heteroscyphus denticulatus
a
Isothecium algarvicum
Isothecium prolixum
a
Jungermannia callithrix
Jungermannia pumila
Lepidozia cupressina
Lepidozia reptans
Leucobryum glaucum
Leucodon treleasei
a
Marsupella adusta
Neckera pumila
Orthotrichum affine
Oxystegus tenuirostris
Pelekium atlanticum
a
Philonotis fontana
Physcomitrium pyriforme
Plagiochila maderensis
a
Plagiothecium succulentum
(continued)
Threatened bryophytes in Madeira and Selvagens Archipelagos 7
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bryophytes list and the Data Deficient taxa (Table 4) but,
as mentioned before, all bryophytes were evaluated and
designated to category.
The Red List
The present bryophyte Red List for the Madeira and Sel-
vagens Archipelagos includes 127 taxa (about 23.6% of
the bryoflora reported), specifically 5 CR (0.9%), 22 EN
(4.1%) and 100 VU (18.6%). From the Madeiran and
Macaronesian endemics, 13 come under a threat category
(57.9%). A relatively high percentage of taxa are found in
the NT category (11.5%) since all endemics that were not
included in a threat category belong to the Near Threat-
ened category, which includes a total of 62 bryophytes.
The number of LC taxa represents about 48.9% of
Madeiran bryoflora (263 bryophytes, including seven con-
sidered LC-att), and 15.2% are included in the DD cate-
gory (including DD-n and DD-va).
Comparing mosses with liverworts we verify a smaller
percentage (21.4%) of threatened mosses and a higher
percentage of threatened liverworts (29.1%) (Table 5).
An analysis of the threatened bryophyte distribution by
habitat (Tables 6 and 7) revealed that ‘high mountain hab-
itats’ comprise the highest number of threatened taxa
(29.5%) and also the highest number of threatened taxa
that are exclusive (48.2%) to the habitat. This is followed
by the Laurel forest habitat where 78.9% of the endemics
also exist, with exactly 39.5% of them exclusive to this
‘relictual’ forest.
On Madeira Island a relatively high frequency of threat-
ened bryophytes is found in certain areas. A significantly
higher number of threatened taxa were found in areas cor-
responding to the Natura 2000 areas (Laurel Forest area
PTMAD0001, Ponta de S~
ao Louren¸co PTMAD0003,
Central Mountain Massif PTMAD0002 and PTZPE0041,
Achadas da Cruz PTMAD0005, Moledos Madalena do
Mar PTMAD0006 and Pin
aculo PTMAD0007 location
on map from EEA, 2013—2014) than in other areas (x
2
¼
26.547, P<0.05). This may be observed in Fig. 1, where
the squares (0.5 km 0.5 km) with higher number of
threatened taxa are represented by larger symbols than
those with a smaller number of threatened bryophytes.
Table 3. (Continued )
Pterigynandrum filiforme
Radula wichurae
a
Rhynchostegiella macilenta
a
Riccardia multifida
Riccia atromarginata
Riccia cavernosa
Riccia ciliata
Riccia crozalsii
Riccia trabutiana
Schistidium apocarpum
Telaranea europaea
Tetrastichium fontanum
Thuidium tamariscinum
Tortula atrovirens
Ulota crispa
a
Macaronesia endemic.
b
Madeira endemic.
c
Under Annex II of the Habitat Directive.
Table 4. Data-deficient (DD) taxa.
Taxa
Mosses
Andreaea rothii;Archidium alternifolium;Barbula convoluta;Barbula unguiculata;Bryoerythrophyllum recurvirostrum;Bryum
muehlenbeckii;Bryum subapiculatum;Bryum ruderale;Bryum sauteri;Campylopus brevipilus;Campylopus cygneus;Campylopus
subulatus;Ceratodon purpureus subsp. stenocarpus;Crossidium squamiferum;Dichodontium flavescens;Dicranella humilis;
Dicranum flagellare;Dicranum montanum;Didymodon acutus;Didymodon australasiae;Entosthodon fascicularis;Entosthodon
muhlenbergii;Ephemerum crassinervium;Fissidens adianthoides;Fissidens bryoides;Fissidens bryoides var. caespitans;Fissidens
crassipes;Fissidens monguillonii;Fissidens rivularis;Fissidens sublimbatus;Fissidens viridulus var. incurvus;Grimmia arenaria;
Grimmia laevigata;Grimmia orbicularis;Grimmia ramondii;Gymnostomum calcareum var. atlanticum;Hygroamblystegium varium
subsp. humile;Isopterygiopsis pulchella;Leptophascum leptophyllum;Mnium hornum;Orthotrichum acuminatum;Orthotrichum
cupulatum;Pelekium minutulum;Philonotis seriata;Plagiomnium ellipticum;Plagiothecium denticulatum;Pohlia bolanderi;Pohlia
melanodon;Pottiopsis caespitosa;Ptychostomum pseudotriquetrum var. bimum;Ptychostomum rubens;Racomitrium affine;
Rhynchostegiella curviseta;Rhynchostegiella trichophylla;Schistidium agassizii;Schistidium rivulare;Syntrichia latifolia;
Syntrichia montana;Syntrichia montana var. calva;Thuidiopsis sparsa;Tortella squarrosa;Tortula acaulon;Tortula bolanderi;
Tortula pallida.
Liverworts and hornworts
Anastrophyllum minutum;Anthoceros agrestis;Calypogeia azorica;Calypogeia azurea Cephalozia catenulata;Cephaloziella
stellulifera;Cheilolejeunea cedercreutzii;Cladopodiella francisci;Conocephalum salebrosum;Jungermannia atrovirens;Leiocolea
heterocolpos;Leiocolea turbinata;Mannia fragrans;Pallavicinia lyellii;Odontoschisma prostratum;Riccardia latifrons;Riccia
bifurca;Riccia ciliifera.
8 M. Sim-Sim et al.
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Additional remarks
Excluded taxa: We were able to exclude seven species
from the Madeiran bryoflora as a result of herbarium revi-
sions, namely: Glyphomitrium daviesii (Dicks.) Brid.,
Gyroweisia tenuis (Schrad. ex Hedw.) Schimp., Leptodon-
tium flexifolium (Dicks.) Hampe, Paraleucobryum longi-
folium (Ehrh. ex Hedw.) Loeske, Pterygoneurum ovatum
(Hedw.) Dixon, Riccardia incurvata Lindb. and Tortella
humilis (Hedw.) Jenn.
Notes on taxa with doubtful taxonomic status:Anoectan-
gium angustifolium Mitt. and Anoectangium aestivum var.
madeirense (Geh.) D€
ull are both now considered as Anoec-
tangium aestivum (Hedw.) Mitt. However, after revising her-
barium material (MADJ 7570; MADJ 5041; MADJ 5042)
we suggest that a taxonomic and molecular study should be
developed to clarify the existence (or not) of distinct taxa.
Amphidium curvipes (M€
ull. Hal.) Broth, once distin-
guished as a Macaronesian endemic species, currently
corresponds to Amphidium tortuosum (Hornsch.) Cufod.
(Ros et al.,2013). However, preliminary results of molec-
ular and taxonomic studies to determine whether this cur-
rent position is correct, indicate that A. curvipes is
actually a separate taxon from A. tortuosum.
Blindia acuta var. madeirensis (Geh.) Perss. was also
removed as an endemic variety and is now considered as a
synonym of Blindia acuta (Hedw.) Bruch & Schimp. Nev-
ertheless, herbarium revisions (MADJ 6593; MADJ 5932;
MADJ 6263; MADJ 5080) throw up a number of doubts
which should be clarified by additional studies.
Homalothecium mandonii (Mitt.) Geh was formerly
recognized as a Macaronesian endemic. Although this
taxon is currently usually considered a synonym of Homa-
lothecium sericeum (Hedw.) Schimp., recent molecular
studies suggest that it deserves recognition at some taxo-
nomic level (D
esamor
eet al.,2012).
Discussion
The present Red List of the bryophyte flora of the
Madeiran and Selvagens Archipelagos was obtained fol-
lowing the IUCN criteria, adapted by Gonz
alez-Mancebo
et al. (2012) to islands. It represents an important step in
the definition of bryophytes (and areas) with conservation
priority and simultaneously increases our knowledge of
the taxa and environments that need further study.
The marked alteration in vegetation composition and
land use in the past 40 years on Madeira Island is a conse-
quence of significant levels of urban and touristic devel-
opment. In fact, touristic development has led to high
levels of pressure in coastal areas. Urbanization, agricul-
ture, hydric resource exploitation and forestry manage-
ment were mainly responsible for the introduction and
expansion of exotic and invasive species in mesic areas,
replacing large areas of natural vegetation. In high altitu-
dinal areas of Madeira, intense unregulated grazing in
areas spanning from the Ericaceous forest and thickets to
the high mountain habitat resulted in degradation of vege-
tation and habitats in several areas. As anthropogenic
activity increased, fire and habitat fragmentation became
more frequent with a clear cumulative impact on natural
vegetation integrity, especially if we consider the island’s
relatively small size.
Consequently, most bryophytes considered as threat-
ened have declined due to the habitat destruction or reduc-
tion that has resulted from agricultural exploitation, urban
construction or habitat modification. Additionally, it is
possible to verify that a large percentage of the bryophytes
Table 5. Comparison between percentage (%) and Number (N)
of liverworts, mosses and hornworts taxa for each category.
Liverworts Mosses Hornworts
%N
%N
%N
CR 0.6 1 1.1 4
EN 4.7 8 3.9 14
VU 4.8 41 16.4 59
Red List 29.1 50 21.4 77
NT 10.5 18 11.9 43 16.7 1
LC 49.4 85 48.1 173 83.3 5
DD 11.1 19 17.5 63
RE or EX 1.1 4
Table 6. Percentage of bryophyte occurrence by habitat for all categories considered.
CR EN VU Red List NT DD LC EX or RE
Coastal habitats/lowland habitats 0.8 11.5 12.3 10.0 16.9 60.8
Mesic areas in the lower parts of the islands 0.8 0.8 8.7 10.3 7.9 10.3 70.3 1.2
High mountain habitats 2.6 5.1 21.8 29.5 10.3 16.7 43.5
Ericaceous forest and thickets (E. patycodon eE. arborea) 1.2 14.8 15.8 10.1 5.1 68.8
Laurel forest 0.3 3.6 18.3 22.2 12.3 9.0 56.2 0.3
Water habitats 1.0 3.1 12.2 16.3 12.2 4.1 67.4
Humanized/disturbed habitats 6.3 87.4 6.3
Threatened bryophytes in Madeira and Selvagens Archipelagos 9
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considered as threatened are currently highly dependent
on the maintenance of the legal protection of their habitats
(Fig. 1), since both the Laurel forest and the Mountain
Massif are covered by national and international legal pro-
tection. However, if current socioeconomic changes result
in a weakening of habitat protection maintenance and hab-
itat surveillance, this will almost certainly increase the
occurrence and magnitude of other threats (e.g. fire
events, invasive species expansion).
For each altitude level there are different threats to be
considered, and threat levels and conservation measures
are therefore considered separately for each major habitat,
taking into account their specific characteristics.
Coastal habitats/lowland habitats
A significant percentage of bryophyte from coastal/low-
land habitats is under threat, which reflects both the loss
of this plant group’s habitat, as a result of construction
and leisure developments, and its highly limited
geographic range (influencing the distribution area). For
instance, Acaulon muticum (Schreb. ex Hedw.) M€
ull. Hal.
and Acaulon triquetrum (Spruce) M€
ull. Hal. are mosses
with an annual life cycle having also a very restricted area
of occurrence (Ponta de S~
ao Louren¸co), and their main
identified threat is ‘Human intrusions & disturbance
Recreational activities (6.1)’ (IUCN, 2012). The Madeiran
endemic liverwort Frullania sergiae Sim-Sim et al. has a
very small geographic range, so we underline the neces-
sity to establish monitoring points to assess population
integrity, especially in Desertas, where the majority of its
populations are to be found, but also in Porto Santo where
human presence is stronger. Similarly, the endemic Riccia
atlantica S
ergio & Perold, a species that grows on poor
soils near cliffs, also has a small geographic range, and
should also be regularly monitored.
The most frequent species to be found in this harsh hab-
itat include, for example, the hornwort Anthoceros agres-
tis Paton, the liverworts Fossombronia caespitiformis De
Not.exRabenh., Lunularia cruciata (L.) Lindb.,
Table 7. Percentage of bryophytes occurring exclusively in one of Madeira’s habitat types by category considered.
CR EN VU Red List NT DD LC EX or RE
Coastal habitats/lowland habitats 2.3 27.9 30.2 16.3 27.9 25.6
Mesic areas in the lower parts of the islands 6.1 6.1 18.1 30.3 3.2 36.3 21.1 9.1
High mountain habitats 7.4 7.4 33.3 48.2 7.4 33.3 11.2
Ericaceous forest and thickets (E. platycodon e and E. arborea)31.8 31.8 4.6 18.2 45.4
Laurel forest 0.7 8.7 28.9 38.5 16.7 13.1 31.2 0.7
Water habitats 14.3 14.3 28.6 71.4
Humanized/disturbed habitats
Fig. 1. Natura 2000 areas (lighter area) and threatened bryophyte distribution (frequency) along Madeira Island.
10 M. Sim-Sim et al.
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Plagiochasma rupestre (J. R. Forst. & G. Forst.) Steph.
and Riccia subbifurca Warnst. ex Croz., and the mosses
Bryum canariense Brid., Eucladium verticillatum (With.)
Bruch & Schimp, Ptychomitrium nigrescens (Kunze)
Wijk & Margad., Tortella flavovirens (Bruch) Broth., Tor-
tula muralis Hedw. and Trichostomum brachydontium
Bruch.
Mesic areas in the lower parts of the islands
This area would probably support a different bryoflora if it
was not as disturbed as it is, since it would potentially
include the Laurisilva Mediterr^
anica do Barbusano natu-
ral forest (Semele androgynaeApollonietum barbujanae
sigmetum). However, some taxa considered as threatened
can be found in remains of natural vegetation, especially
close to streams in narrow valleys. For instance, according
to Lobo et al. (2011), the moss Fissidens ovatifolius R.
Ruthe is currently highly threatened due to human activi-
ties as well as landslides and torrents. In this general habi-
tat type, the most widespread (LC) species that can be
found, considering also disturbed sites and exotic forest,
are Bartramia stricta Brid., Dialytrichia mucronata
(Brid.) Broth., H. sericeum,Kindbergia praelonga
(Hedw.) Ochyra, Oxyrrhynchium hians (Hedw.) Loeske,
T. brachydontium and Timmiella barbuloides (Brid.)
M€
onk, among the mosses, and Fossombronia angulosa
(Dicks.) Raddi, Frullania azorica Sim-Sim et al., Hetero-
scyphus denticulatus (Mitt.) Schiffn., Marchantia poly-
morpha L., Riccardia chamedryfolia (With.) Grolle and
Targionia hypophylla L. among the liverworts.
Laurel forest
Despite the fact that the best-preserved area of laurel for-
est in Macaronesia is to be found on Madeira Island, there
are certain threats that need to be taken into consideration,
such as the impact of touristic activity and exotic species
invasion. In fact, several areas of Laurel forest have been
disturbed, which justifies the need for active management
to promote habitat restoration.
A high percentage of endemic (c. 39%) and threatened
bryophytes (c. 38%) are exclusive to the Laurel forest,
which indicates the importance of maintaining the habitat
diversity and conservation policies that already exist. In
fact, as shown in Fig. 1, there are some areas in the Natura
2000 habitats that should be highlighted, given the high
concentration of threatened taxa.
Aphanolejeunea azorica (V. Allorge & Ast) Bernecker
&P
ocs, Cololejeunea schaeferi Grolle, Lejeunea canar-
iensis (Steph.) Steph., Porella inaequalis (Gottsche ex
Steph.) Perss., Tylimanthus madeirensis Grolle & Perss.
or Echinodium spinosum (Mitt.) Jur. (the latter under
Annex II of the Habitat Directive) are all examples of
threatened endemic species, currently restricted to the
best-preserved areas of the Laurel forest.
On the other hand, the most frequent and dominant taxa
occurring in a typical Laurel forest stand are Andoa ber-
thelothiana (Mont.) Ochyra, Fissidens serrulatus Brid.,
Hypnum cupressiforme Hedw., Isothecium prolixum
(Mitt.) M. Stech, Sim-Sim, Tangney & D. Quandt, Leuco-
bryum juniperoideum (Brid.) M€
ull. Hal. and Tetrastichium
virens (Cardot) S.P. Churchill among the mosses, and
Calypogeia fissa (L.) Raddi, Conocephalum conicum (L.)
Dumort., Diplophyllum albicans (L.) Dumort., Drepano-
lejeunea hamatifolia (Hook.) Schiffn., Frullania spp. (F.
teneriffae (F. Weber) Nees, F. tamarisci (L.) Dumort., F.
microphylla (Gottsche) Pearson), Harpalejeunea molleri
(Steph.) Grolle, Lejeunea spp. (e.g. L. lamacerina (Steph.)
Schiffn., L. flava subsp. moorei (Lindb.) R. M. Schust., L.
eckloniana Lindenb.), Lophocolea bidentata (L.)
Dumort., Metzgeria furcata (L.) Dumort., Microlejeunea
ulicina (Taylor) A. Evans, Plagiochila bifaria (Sw.) Lin-
denb., Plagiochila stricta Lindenb., Porella canariensis
(F. Weber) Underw., Radula carringtonii J. B. Jack, Rad-
ula lindenbergiana Gottsche ex C. Hartm. and Saccogyna
viticulosa (L.) Dumort. among the liverworts.
Ericaceous forest and thickets
In this habitat type, we find some exclusive threatened spe-
cies such as Plagiochila spinulosa (Dicks.) Dumort., and
the moss Rhytidiadelphus loreus Hedw.) Warst., occurring
especially on the borders of the ericaceous forest. The
destruction of vascular plant vegetation through fires and
grazing has historically been one of the main threats affect-
ing these habitats, and has resulted in a decline in ecosystem
quality and bryophyte populations, which were dependent
on that vegetation for shelter (S
ergio et al.,2008). Compar-
ing the Ericaceous forest and thickets communities with the
Laurel forest, it is possible to observe a difference in the
bryoflora composition and dominance with particular refer-
ence to the Scapanio gracilisDicranetum scottiani com-
munity (Fontinha et al.,2006).
The most common bryophyte community includes,
besides Scapania gracilis Lindb. and Dicranum canar-
iense Hampe ex M€
ull. Hal., the mosses Atrichum undula-
tum (Hedw.) P. Beauv., H. cupressiforme,Pogonatum
aloides (Hedw.) P. Beauv., Polytrichum commune Hedw.,
P. juniperinum and Pseudoscleropodium purum (Hedw.)
M. Fleisch., and the liverworts Drepanolejeunea hamati-
folia (Hook.) Schiffn., F. teneriffae,L. lamacerina,M. uli-
cina,Plagiochila exigua (Taylor) Taylor and Plagiochila
punctata (Taylor) Taylor.
High-mountain habitats
In the Madeiran Archipelago this type of ecosystem is
restricted to a small area of the main island, and the
Threatened bryophytes in Madeira and Selvagens Archipelagos 11
Downloaded by [Susana Fontinha] at 10:07 29 May 2014
percentage of threatened taxa that are exclusive to this
environment in Madeira is the highest (48.2%). This can
be explained in part by the small area present on Madeira
Island, but also by the fires and grazing that have histori-
cally affected these areas in recent decades, causing a sig-
nificant destruction and decline in the bryophytes’
extension.
For example, species like Andreaea flexuosa subsp. lui-
sieri (S
ergio & Sim-Sim, 2012), A. heinemannii Hampe &
M€
ull.Hal. subsp. heinemannii,A. rupestris Hedw., Anaco-
lia webbi (Mont.) Schimp. and Schistidium strictum
(Turner) Loeske ex are to be found on the main island
mainly above 1500 m, and therefore have limited natural
distribution on Madeira. Consequently, anthropogenic
influences on Madeira’s high-altitude ecosystems, and
projections of the effects of climate change, represent
challenges to these species’ survival.
The commonest bryophytes present are limited to a
small group of taxa, the most prominent being Amphidium
mougeotii (Bruch & Schimp.) Schimp., Andreaea spp.,
Antitrichia curtipendula (Hedw.) Brid., Campylopus
pilifer Brid.,Grimmia spp., I. prolixum,Polytrichum pili-
ferum Hedw., Ptychomitrium polyphyllum (Dicks. ex Sw.)
Bruch & Schimp. and Racomitrium spp. among the
mosses, and F. tamarisci,Marsupella emarginata (Ehrh.)
Dumort. and Scapania compacta (A. Roth) Dumort.
among the liverworts.
Water habitats
Some important endemic species such as Bryoxiphium
madeirense
A. L€
ove & D. L€
ove (Madeiran endemic) are
highly dependent on the maintenance of water habitat
quality and quantity, and have a very restricted habitat
(vertical slopes with dripping water). In 2010 a wildfire
reached one of the largest B. madeirense populations,
which was reduced to less than half of its original num-
bers. Another Madeiran endemic, Thamnobryum fernan-
desii S
ergio also grows in permanently wet habitats (rocks
or waterfalls from sea level up to 1000 m in the central
part of Madeira Island, where it is more common). Rham-
phidium purpuratum Mitt. is currently in the Near Threat-
ened category; however, since it is considered Vulnerable
in the Canary Islands (Gonz
alez-Mancebo et al.,2012)
and in Europe (ECCB, 1995) it is necessary to pay atten-
tion to this species’ populations.
With regard to Madeira Island, the latest governmental
reports demonstrate a trend of reduced water availability.
Projections indicate a significant reduction in the volume
of water annually available for recharge and runoff of
about 30% by 2050 and 40% by the end of the century.
These projections represent a threat that is anything but
negligible, especially if we consider sensitive bryophytes
that already have a small distribution area, occurring in
restricted habitats such as small waterfalls (Lu
ıs et al.,
2012).
We generally find the following dominant bryophyte
species: Atrichum androgynum (M€
ull. Hal.) A. Jaeger,
Brachythecium rivulare Schimp., Ptychostomum pseudo-
triquetrum (Hedw.) J.R. Spence & H.P. Ramsay, Didymo-
don insulanus (De Not.) M.O. Hill, Fissidens spp. (F.
sublineaefolius (P. de la Varde) Brugg.-Nann., F. asple-
nioides Hedw., Fissidens taxifolius subsp. pallidicaulis
(Mitt.) M€
onk.), Fontinalis antipyretica Hedw., Hygroam-
blystegium fluviatile (Hedw.) Loeske, Leptodictyum ripa-
rium (Hedw.) Warnst., Philonotis rigida Brid.,
Plagiomnium undulatum (Hedw.) T.J. Kop., Plagiomnium
undulatum var. madeirense T.J.Kop. & S
ergio, Rhynchos-
tegium riparioides (Hedw.) Cardot, Rhizomnium puncta-
tum (Hedw.) T.J. Kop. and Thamnobryum alopecurum
(Hedw.) Gangulee among the mosses, and L. cruciata,
Scapania undulata (L.) Dumort. and C. conicum among
the liverworts.
We would underline the consistency with results
obtained by Schnyder et al. (2004), Natcheva et al.
(2006), S
ergio et al. (2007,2013), Papp et al. (2010) and
Gonz
alez-Mancebo et al. (2012) showing that liverworts
display a higher percentage of Red Listed taxa compared
with mosses (Table 5). This can be explained by the
importance of the ecological and climatic conditions of
the laurel forest, which must therefore be retained in the
interests of successful bryoflora preservation, as was also
concluded by Gonz
alez-Mancebo et al. (2012).
In fact, several studies highlight the similarities
between the bryoflora of the Canary Islands, Madeira and
Selvagens archipelagos. For example, the bryovegetation
of arid and semi-arid environments reveals high levels of
floristic similarity between the driest islands of the
Madeira region (Selvagens, Desertas and Porto Santo
islands) and Fuerteventura and Lanzarote (Canary Islands)
(Sim-Sim et al., 2010). For both Red Lists, these habitats
have a lower percentage of threatened taxa, even consider-
ing the occurrence of particular endemic taxa.
Additionally, at the community assemblage level, there
is a high level of similarity between the Canary Islands
and Madeiran epiphytes (Zippel, 1998,K
€
urschner,
2007a), such as the alliance EchinodioNeckerion inter-
mediate, typical of the Laurel forest. In this habitat it is
possible to verify that Red Listed taxa exist that are com-
mon to both the Canary Islands and Madeira, for instance,
Echinodium spinosum and Radula jonesii (despite being
in different threat categories).
Conclusions
In Madeira, climatic factors such as rainfall and tempera-
ture, related to exposure and altitude, are the major
factors influencing bryophyte distribution. However,
12 M. Sim-Sim et al.
Downloaded by [Susana Fontinha] at 10:07 29 May 2014
micro-environmental features (pH, shade, air humidity,
water proximity) are decisive in the micro-community
and for the stability of threatened bryophyte populations
(Hedderson & Brassard, 1990). A large number of the cur-
rent Red List taxa are highly dependent on the habitat
micro-conditions, and a significant number of them have
been severely affected by the impacts of the overstocking
of sheep and goats, the canalization of watercourses or
vascular plant replacement. Furthermore, some of the Red
Listed populations are so small that they might easily be
wiped out by single events (e.g. F. sergiae and B.
madeirense).
Bryophytes under the NT and DD categories deserve
particular attention despite the fact that they are not con-
sidered as under threat, as they may be moved up into a
threat category when more information has been acquired
(DD), or if their habitats change drastically (NT) (Natch-
eva et al.,2006). For instance, in high-mountain habitats,
the high percentage of threatened bryophytes that are
exclusive to this ecosystem, plus the high percentage of
Data Deficient (c. 33%) cases, points to the urgent need
for monitoring of these populations.
The flora of small islands generally have limited possi-
bilities to migrate horizontally (Montmollin & Strahm,
2005), which increases the risk for species extinction and
reinforces the need to implement effective conservation
measures.
Bryophyte conservation is undoubtedly crucial, since
this taxonomic group plays a fundamental role in ecosys-
tem functioning, and is involved in important large-scale
carbon and nutrient cycling, including nitrogen-fixing in
association with cyanobacteria (Chapin et al.,2000;
Beringer et al.,2001; Lindo et al.,2013). They form dense
carpets in several ecosystems, influencing numerous eco-
logical processes and community assembly (Schofield,
2000). Bryophytes’ economic value is still underesti-
mated, since in many regions (including the Madeiran
autonomous region) they are not considered as an essen-
tial part of the natural capital. An important step to
address this issue would be to evaluate the ecological
services provided by bryophytes in different habitats and
define Important Bryophyte Conservation Areas (IBrA)
based on Red List results and criteria applied to bryo-
phytes (e.g Papp, 2008; Lockhart et al.,2012).
An important result of the identification of hotspots for
bryophytes, especially those based on Red Listed taxa, is
the definition of reserves or microreserves, with specific
habitat conservation initiatives as well as specific manage-
ment guidelines for the sustainable use of the areas
(Rodrigues et al.,2006). This information is also relevant
insofar as it interlinks with data from other taxonomic
groups and can be used for the Red List Index
development.
As an example, one priority species is Marsupella pro-
funda Lindb., one of the three Annex II priority species to
be found in Madeira and inhabiting acid, clay-rich,
exposed soils. This worldwide threatened species requires
Special Areas of Conservation. In protected areas, threats
can be prevented or minimized, by increasing land manag-
ers’ knowledge of the presence of threatened plants and by
implementing monitoring systems to improve the moni-
toring of the development of populations.
Species threat status evaluation is a continuous process,
as new data-gathering or taxonomic revisions may provide
information that leads to changes in the number of taxa
within a category (Bilz et al.,2011). Moreover, there is
always a high level of uncertainty in ecological processes,
since knowledge of cumulative effects and relations
between different taxonomic groups is still under study
and development.
In conclusion, for such a small area of territory, the
Madeira archipelago (Madeira Island in particular) boasts
an extremely diverse bryoflora. Compared with the Portu-
guese mainland and the Canary Islands (Macaronesian
islands), which are also regions with high levels of diver-
sity, but with considerably larger areas, Madeira stands
out as an important bryophyte diversity hotspot.
Acknowledgements
We would like to express our gratitude to Juana Mar
ıa
Gonz
alez-Mancebo for her elucidation of the IUCN crite-
ria. We would also like to thank Leena Lu
ıs for all the
work developed over the past years, which provided
important information on Madeira bryoflora. We are also
grateful to Tereza Almeida for her help with georeferenc-
ing and to F
abio Reis for his help with fieldwork. This
work was supported by FCT (Funda¸c~
ao para a Ci^
encia e
Tecnologia) under Grant no. PTDC/AGR-CFL/111241/
2009.
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