Global moss diversity: spatial and taxonomic

patterns of species richness

Jan Laurens Geffert, Jan-Peter Frahm, Wilhelm Barthlott, Jens Mutke
Nees-Institut fur Biodiversitat der Pflanzen, Rheinische Friedrich-Wilhelms-Universitat Bonn, Germany
We have analysed the global patterns of moss species diversity based on a dataset created from checklists,
online databases, and herbarium records. We collected more than 100 000 distribution records for over 400
different geographical units and standardized species taxonomy using the TROPICOS database of the
Missouri Botanical Garden. Maps of overall moss species richness, as well as individual maps for taxonomic
orders of mosses, are provided. Based on our dataset, we did not find a general latitudinal gradient of
increasing moss diversity with decreasing latitude. Several areas of temperate broadleaf forests, boreal
forests, and tundra show relatively high species richness that is comparable to tropical regions. Centres of
moss diversity include the northern Andes, Southeast Asia, Mexico, and Japan, as well as the Himalayan
region, Madagascar, the East African Highlands, central Europe, Scandinavia, and British Columbia. Our
dataset presents the first collection of moss species inventories with global coverage. It contributes to
documentation and understanding of global biogeographic patterns in mosses, helps to identify gaps in
floristic knowledge, and could prove to be a valuable resource to aid taxonomic and systematic revisions or
assessments of species and genera, by quickly and easily supplying an overview of the geographic
distribution of a given taxon.
Keywords: Biogeography, Bryophyte diversity, Latitudinal gradient, Macroecology, Moss species richness

Introduction

2010). They can be the determining element of

vegetation in bog or tundra ecosystems (Vitt &
Wieder, 2008), and are key species of primary
succession (Goffinet & Shaw, 2008), water retention
(Churchill, 2009), carbon sequestration (Hallingback
& Hodgetts, 2000; de Lucia et al., 2003), nutrient
cycling (Chapin et al., 1987; Turetsky, 2003), and soil
erosion control (Hallingback & Hodgetts, 2000).
Some documentation of the observed patterns in
bryophyte species richness does exist. Von Konrat
et al. (2008) used floristic inventories of 400 regions
worldwide to map the worlds species richness in
liverworts, and Villarreal et al. (2010) summarized the
global distribution of hornworts. But although (1)
substantial progress has been made towards cataloguing the worlds moss species (e.g. Crosby et al., 1999;
Magill, 2010; Royal Botanic Gardens Kew, 2010), (2)
the available data on mosses is generally believed to be
more reliable than that on e.g. liverworts (von Konrat
et al., 2008), and (3) sampling intensity and cataloguing completeness in many tropical countries have
increased (Shaw et al., 2005), actual analyses on
continental or global scales are still sparse.

Species diversity is unequally distributed across the

globe (Barthlott et al., 2007). Exploration and documentation of the spatial patterns of species richness
are of crucial importance for the conservation and
sustainable use of biological resources (Mutke &
Barthlott, 2005). Maps of species diversity allow for
the concise and efficient display of these spatial patterns
and can be an important tool to visualize information,
present findings, and inform policy makers or the
general public. What is more, understanding the
underlying mechanisms constitutes one of the most
significant intellectual challenges to ecologists and
biogeographers (Gaston, 2000).
During the last decades, large-scale patterns of
species diversity have been increasingly documented
for various groups of organisms. In plants, detailed
global diversity maps for vascular plants (Barthlott
et al., 1996; Barthlott et al., 2007; Kreft & Jetz, 2007),
gymnosperms (Mutke & Barthlott, 2005), ferns and
fern allies (Kreft et al., 2010), and even benthic
marine algae (Kerswell, 2006) are now available.
Bryophytes, in particular mosses, are an important
element of land plant diversity (Hallingback & Tan,

Conservation of bryophyte diversity

Correspondence to: Jan Laurens Geffert, Nees-Institut fur Biodiversitat der

Pflanzen, Rheinische Friedrich-Wilhelms-Universitat Bonn, Meckenheimer
Allee 170, 53115 Bonn, Germany. Email: geffert@uni-bonn.de

Biodiversity is rapidly declining across the globe

(Barnosky et al., 2011). This is increasingly raising
concerns amongst scientists and policy makers, and

British Bryological Society 2013

DOI 10.1179/1743282012Y.0000000038

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Global moss diversity

has led to a variety of efforts to combat loss of

biodiversity (Balmford et al., 2005), among them the
Convention on Biological Diversity and the Global
Strategy for Plant Conservation (GSPC).
The GSPC set 16 targets for the year 2010 to slow
the pace of plant extinction worldwide. Target 1 was to
establish a working list of all plant species, including
bryophytes, and has been met through the creation of
The Plant List (Royal Botanic Gardens Kew, 2010), a
joint project of Convention on Biological Diversity,
the Royal Botanic Gardens Kew, Missouri Botanical
Garden and several other partners.
However, identifying all relevant species can only be
one step towards the conservation of global moss
diversity. Target 5 of the GSPC was to assure the
protection of 50% of the most important areas for plant
diversity, but little work has been done to identify areas
of particular relevance for mosses. These data are often
available, but widely scattered throughout the relevant
literature (compare von Konrat et al., 2010).

Centres of moss diversity

So far, most delineations of centres of moss diversity
in textbooks rely on a mix of data collection and
expert opinion (Bates & Farmer, 1992; Goffinet &
Shaw, 2008; Shaw & Goffinet, 2000). This approach
is distinguished by little methodological transparency
and influenced by personal opinion. For example,
Tan and Pocs (2000) produced a list of moss diversity
hotspots per continent, but did not differentiate
between centres of species richness and centres of
endemism. As a result, species rich regions like the
northern Andes were listed side by side with regions
of relatively low species richness but distinct species
inventories, like the Mediterranean.
We believe that it is important to disentangle these
different dimensions of diversity and to assess species
diversity in a numerical and reproducible way. First
work of this type on mosses was started by our working
group in 1998, with a preliminary dataset published in
Mutke and Barthlott (2005). The resulting map, however, was still coarse in resolution and did not achieve
global coverage. Furthermore, this dataset consisted of
simple documented species numbers instead of entire
species lists, which increased taxonomic inflation and
caused a bias towards well collected but badly revised
regions or checklists that included any recorded species
name regardless of its taxonomic status.
For this paper, we used national and regional species
checklists as well as online databases and herbarium
records to assess the global patterns of moss diversity in
a quantitative macroecological approach.

Methods
Data collection
We collected moss species inventories of political or
regional units worldwide (hereafter geographical

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units, GUs) from the literature and merged them as

species lists into a single dataset (inventory-based
diversity mapping approach, compare Barthlott
et al., 1999, and Mutke & Barthlott, 2005). Main
data sources were national and regional checklists,
floras, and digital online databases (Table 1). We
sought to achieve global coverage, but included only
non overlapping GUs. If multiple data sources were
available for a given region, we chose the most recent
publication that was available to us.
Table 1 Data sources
Region
Antarctica
Argentina
Australia
Austria
Bangladesh
Belgium
Bhutan
Bolivia
Brazil
Canada
Central and
South America
Central Europe
Chile
China
Corsica
Estonia
Former Soviet Union
France
Galapagos

Source
Ochyra et al., 2008
Matteri, 2003
Klazenga et al., 2009
Kockinger et al., 2008
OShea, 2003a
Sotiaux et al., 2007b
Long, 1994
Churchill et al., 2009
Forzza et al., 2010
Ireland et al., 1987
Delgadillo et al., 1995

Hill et al., 2006

Muller, 2009
Redfearn et al., 1994
Sotiaux et al., 2007a
Ingerpuu et al., 1994
Ignatov et al., 2006
Lemonnier, 2010
Ziemmeck and
Gradstein, 2009
Germany
Koperski et al., 2000
Great Britain
Hill et al., 2008
Greenland
Goldberg, 2003
Guianas
Boggan et al., 1997
Hawaii
Staples et al., 2004
Hong Kong
Hu et al., 2002
Hungary
Erzberger and Papp, 2004
Iceland
Johannsson, 2003
India
Lal, 2005
Indochina
Tan and Iwatsuki, 1993
Japan
Iwatsuki, 2004
Korea
Park and Choi, 2007
Liechtenstein
Senn, 2000
Luxembourg
Werner, 2008
Malesia
Tan and Soon, 1997
Middle East
Frey and Kurschner, 1991
Mongolia
Tsegmed, 2001
Netherlands
Siebel et al., 2005
New Caledonia
Pursell and Reese, 1982
New Zealand
Fife, 1995
North Africa
Ros et al., 1999
Pakistan
Higuchi and Nishimura, 2003
Seychelles
Frahm and Ho, 2009
Slovakia
Kubinska and Janovicova, 1998
South Africa
Germishuizen and Meyer, 2003
Southeast Europe
Sabovljevic et al., 2008
Sri Lanka
OShea, 2002
Sub Saharan Africa
OShea, 2003b
Sulawesi
Gradstein et al., 2005
Sumatra
Ho et al., 2006
Svalbard
Frisvoll and Elvebakk, 1996
Switzerland
Nationales Inventar der
Schweizer Moosflora, 2010
Taiwan
Chiang et al., 2001
Thailand
He, 1997
United States of America The New York Botanical Garden, 2003

Geffert et al.

Several sources listed subsets for administrative

units or geographical regions of the covered area, and
these subsets were also integrated where available.
However, some subunits were too small scale for our
purpose (e.g. counties of the UK in Hill et al., 2008),
or showed obvious lack of collection intensity (e.g.
Argentine provinces in Matteri, 2003), and were thus
aggregated to larger units.
For most federal states of the USA, no up to date
species list had been published and data from the
Bryophyte Flora of North America were not yet
available (pers. comm., Zander, 2010). We therefore
generated preliminary species lists from records of the
New York Botanical Garden Virtual Herbarium
American Bryophyte Catalog (The New York
Botanical Garden, 2003), by downloading all records
and grouping them by state and taxon.

Taxonomic standardization
Because the data were derived from diverse sources,
the quality of individual lists varied with age, scope,
and taxonomic concept of the original publication.
Older checklists listed outdated taxon names and
occasionally several synonymous names at a time.
Checklists varied in the way they were compiled (see
review of liverwort checklists in Soderstrom et al.,
2008), being based on literature records, verified
specimens, or field collections. Finally, different
authors adhered to different taxonomic concepts,
splitting or lumping different taxa, potentially compromising the comparability of species lists from
different sources by inflating or deflating species
numbers, and thus distorting observed patterns (as
was shown by Isaac et al., 2004).
We therefore deemed it important to standardize
our dataset using a reference species list. The
TROPICOS database of Missouri Botanical Garden
proved to be a unique resource for this purpose. This
database is freely available, continuously updated, and
includes the data of the Crosby et al. (1999) world
checklist of mosses (see Magill, 2010 for an in-depth
discussion). It has been frequently used as the standard
reference in the past and became the sole input for
mosses into The Plant List, the working list of all
known plant species (Royal Botanic Gardens Kew,
2010).
We linked our dataset to TROPICOS and extracted
further information on each given taxon in an
automated procedure: first, we used the automated
name matching tool of the TROPICOS website (http://
www.tropicos.org/NameMatching.aspx) to obtain the
unique identification number of each taxon used by the
TROPICOS data base system. This identification
number enabled us to access more detailed information
with R statistical software (R Development Core Team,
2011). The basic mechanism is to generate a specific web

Global moss diversity

address for every given taxon and query the application

programming interface service to retrieve the desired
information from the database in Extensible Markup
Language or JavaScript Object Notation format (a
similar approach is discussed in Carvalho et al., 2010,
and excellent documentation as well as corresponding
R-scripts can be found on http://www.plantminer.org).
Among the retrieved information was the nomenclatural status of each given taxon (accepted/no
opinion/invalid/illegitimate), as well as an accepted
name for many invalid or illegitimate taxa. By
reassigning these accepted names to invalid and
illegitimate taxa, we standardized our dataset to a
uniform taxonomic concept.

Mapping of species richness

For quantitative analysis of our dataset, the standardized species lists were aggregated by region to
generate species numbers. Taxa listed as illegitimate
and invalid were excluded from the analysis. We
created a map of all GUs included in our dataset
using ArcGIS (ESRI, 2009). For the visualization, a
circle of 200 km radius was added for small oceanic
islands and archipelagos to increase visibility in the
maps. We joined the information on documented
species richness to the corresponding GUs, using the
alpha-2 Codes for the representation of names of
countries (ISO 3166), expanding these where necessary to code for sub national regions. Grades of
documented species richness were visualized in the
map using shades of grey (Figure 1).
The GUs used in this analysis varied widely in size.
We calculated the area of each region in ArcGIS and
used the species area model of Arrhenius (1921),
to estimate species richness per standard area
(Figure 2), using the formula:
 z
A1
S1 ~S2
A2
where S1 is the standardized species richness (per
standard area), S2 is the documented species richness
for the given region, A1 is the standard area (here:
100 000 km2), A2 is the area of the given region and z
is the index of floristic heterogeneity.
This approach uses the observed values (A2, S2) to
calculate the theoretical number of species (S1) for a
theoretical region of the standard area (A1), based on
the observed floristic heterogeneity (z) in the species
inventories (for a detailed comparison of different
species area methodologies refer to Kier et al., 2005).
Based on earlier analyses of the dataset (Mutke &
Geffert, 2010), a z factor of 0.15 was used for this
operation.

Mapping of major subgroups

Collecting species lists instead of species numbers
made it possible not only to analyse moss species

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Global moss diversity

Figure 1 Documented patterns of global species richness in mosses based on national and regional checklists. Numbers
include all taxa listed as legitimate or no opinion by TROPICOS, but exclude illegitimate and invalid names that could not
be resolved.

diversity, but also to map any subset of the data

equivalently. Using the TROPICOS information on
the species families, we linked each species to one of
the major lineages of mosses (Orders). We then
mapped the subgroups of mosses by displaying species

number per region per group. Classification was based

on Syllabus of Plant Families. Part 3: Bryophytes and
Seedless Vascular Plants (Frey et al., 2009). We also
assessed the composition of floristic kingdoms by
aggregating the data for GUs and plotting the number

Figure 2 Patterns of global species richness in mosses. Species numbers have been standardized for an area of 100 000 km2
using the species area model of Arrhenius (1921).

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Global moss diversity

Figure 3 Major orders of mosses for the main floristic kingdoms. The size of the pie charts indicates the total species richness
for the respective region. Global plot shown in the bottom left and regional plots for floristic kingdoms shown in the map.
Delineation of floristic kingdoms was adopted from Olson et al. (2001), but modified by merging Nearctic and Palaearctic regions.

of species per order in pie charts (Figure 3). Floristic

kingdoms were adopted from the biogeographic realms
Olson et al. (2001), but modified by merging Nearctic
and Palaearctic regions.

Results
Species inventories and taxonomic
standardization
Inventory data covering the entire continental global
land surface were collected from a total of 55 national
and regional checklists, floras, and online databases
(listed in Table 1) on over 400 GUs. The final dataset
had over 100 000 records of moss species occurring in
a particular GU.
Of the original 14 804 species in the dataset, 13 371
were automatically matched to a TROPICOS entry as
our taxonomic reference list, 1362 species had to be
manually corrected because of typing errors, aberrant
authority abbreviations and superfluous or missing
hyphens, and 71 names could not be referenced to a
TROPICOS entry and were thus omitted in the further
analysis.
With the taxonomic standardization, 2327 species
names were reassigned to a new taxon name and the
total number of species in our dataset was reduced to
11 388 species. Of these, the vast majority (9272) were
listed as legitimate in TROPICOS. No current
opinion was available for 1925 taxa, while 173 names
were listed as invalid and 18 as illegitimate with no
accepted name to reassign them to.

Mapping of species richness

Moss species were recorded from almost every single
region worldwide, but species richness was unevenly
distributed (Figure 1). Highest documented species
richness was found in the northern Andes, Southeast
Asia, Mexico, and Japan, followed by the Himalayan
region, Madagascar, the East African Highlands, as
well as central Europe, Scandinavia, and British
Columbia. Low documented species richness was
found for deserts and arid regions of the Sahara,
Arabian Peninsula, Namib, Atacama, Caatinga,
parts of Australia, and central Asia.
Areas of temperate broadleaf forests, boreal forests,
and tundra showed relatively high species richness,
with Siberia, Canada, Greenland, Tasmania, and
Tierra del Fuego having equally numerous inventories
as for example the Brazilian Atlantic Forest or central
Africa. The only GU without any recorded moss
species in our dataset was Western Sahara. The
Greenland ice sheet and continental Antarctica were
excluded from the maps as species records were almost
exclusively from coastal areas.

Distribution and diversity patterns of major

subgroups
The individual subgroups showed extremely variable
distribution patterns (Figure 4).
For larger groups like Hypnales, Bryales, and
Dicranales, overall patterns are similar to the total
richness of bryophytes, with members in almost every
single region and high richness in the global centres of

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Figure 4 Patterns of species richness for the major subgroups of mosses. Species numbers as species per region, not
standardized by area. Classification based on TROPICOS and the Syllabus of plant families. Part 3: Bryophytes and seedless
vascular plants (Frey et al., 2009).

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Figure 4b

species diversity. The Hypnales show increased

richness in Southeast Asia and the Himalayas, while
the northern Andes are less pronounced for this
group. The Bryales are especially species rich in
temperate, boreal regions, but Himalaya, Andes, and
East African Highlands show high species richness,
too. The Dicranales show a higher species richness in

the tropics, with the highest values in the East

African Highlands, Himalaya, and Australasia, but
they are also diverse in the Andes, the Amazon Basin
and southern India.
Hookeriales, Hypnodendrales, and Hedwigiales are
examples of primarily tropical groups. The Hookeriales
extend to coastal oceanic areas of the temperate zones

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with just a few species. With over 20 species per region,

Southern Chile, New Zealand, and Tasmania show
higher values than areas of the northern hemisphere.
The Brazilian Atlantic Coast Forest, Central Africa,
East African Highlands, and Queensland also show
relatively high values, but highest species richness of the
Hookeriales is found in the northern Andes, Mexico
and Australasia. The Hypnodendrales is pan-tropical,
although for large parts of the Neotropics, Racopilum
intermedium Hampe is the only species. In southern
Chile, additional Racopilum species increase the diversity of the subgroup. Centres of Diversity are New
Zealand, Australasia, the East African Highlands,
Madagascar, and West Africa. The Hedwigiales have
a Neotropical-African disjunct distribution. The genera
of Hedwigia P.Beauv. and Braunia Bruch & Schimp.
occur in the majority of the northern hemisphere, but
high species richness is mainly found in Peru, Ecuador,
and Bolivia, as well as the East African Highlands.
Interestingly, the group is almost absent from
Australasia and Melanesia.
Smaller groups are much more confined in their
geographical distribution. The Rhizogoniales are pantropical and southern hemispheric with the majority of the
species in eastern Australia, Southeast Asia, New
Zealand, and New Caledonia. Two species occur in
Japan and a single species, Leptotheca gaudichaudii
Schwagr. was reported from Ireland, but according to
Fisk (2008), it is a naturalized epiphyte introduced on
imported tree ferns. Timmiidae occur across the
Holarctic, mainly in boreal regions. There is a disjunct
population of Timmia norvegica J.E.Zetterst. on the
southern island of New Zealand, which is believed to stem
from a natural long range dispersal event (Horton, 1983).
The Andreaeopsida are patchily distributed all over the
world. They are absent from extremely dry desert and
continental regions as well as the lowland tropical forests,
but occur in the boreal zone and all major mountain
ranges. Highest diversity is found in southern Chile,
Tierra del Fuego, New South Wales, and Tasmania.

Discussion
The data presented here give a much more detailed
picture of the global distribution of moss species
richness than any comparable prior analysis.
The fact that mosses occur in high numbers from
polar to tropical regions illustrates their high adaptability to different environmental conditions. The low
species richness in many dry-land regions, however,
demonstrates their high dependence on freely available water in the form of precipitation or cloud
moisture. Only a few specialized taxa manage to
reduce this dependence on moisture and are thus able
to survive in dry-land areas (Frahm, 2001).
The use of data derived from checklists and floras
is not without problems (Kreft et al., 2010), but

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proved to be the most viable approach for mosses,

given the lack of available specimen data or range
maps for many taxa. Some of the species richness
patterns documented here were quite foreseeable and
most likely reflect a real world biogeographic pattern
(e.g. dry regions being very species poor). Others are
most obviously artefacts of uneven bryological
exploration (e.g. extremely low species numbers in
some African countries, compare Mutke & Geffert,
2010).

Latitudinal gradient of species richness

Our findings of equally diverse regions in temperate
and tropical regions seem to conflict with the paradigm
of high diversity in the tropics, as well as previous
reports stating that about two-thirds of all moss
species are tropical species (Frahm, 2003; Gradstein,
2001; Tan & Iwatsuki, 1996; Tan & Pocs, 2000). The
absence of a latitudinal gradient in moss species
richness could be a pure artefact of biased sampling
intensity. Arguably, moss inventories of tropical
regions are far from complete (compare study by
Bebber et al., 2007, for Southeast Asia). Most
temperate regions, particularly Europe, have been
inventoried much more thoroughly than comparable
tropical regions and the documented patterns are
therefore definitively impaired.
But at the same time, taxonomic revisions for
many tropical taxa are still missing (Magill, 2010). It
was stated that the countering effects of increased
sampling and increased synonymy tend to result in
relatively constant species numbers for these regions
(Hedenas, 2007). This is illustrated by the history of
moss checklists from Brazil. Yano (1981) listed 1896
moss species for Brazil. This list was updated several
times and new national species records were added.
However, after revision of taxonomy, specimen and
literature records, Costa et al. (2010) ended up with a
list of only 885 species for the entire country.
Some authors also pointed out that tropical
lowland forests are relatively species poor, and high
species numbers were only achieved in the mountain
ranges and cloud forest ecosystems. Churchill (2009)
stated that the elevational belts of the Andes were the
only reason not to see mosses as an exception from
the latitudinal gradient of species richness, with
temperate taxa in high altitudes contributing significantly to the overall regional diversity on a
macroecological scale.
In contrast, von Konrat et al. (2008) found a
significant latitudinal gradient of species richness for
global liverwort diversity. With a similar degree of
floristic knowledge for mosses and liverworts, it is
difficult to tell why one group would display a strong
bias in geographical sampling while the other does
not. In the tropical zone, Lejeuneaceae contribute

Geffert et al.

heavily to the species diversity of liverworts (von

Konrat et al., 2008). It seems likely that the key
innovation was the adaptation to the epiphyllous
habitat, which is dominated by this group. In
contrast, this habitat is characterized by only very
few mosses.
Several other recent studies focusing on the issue of
a latitudinal gradient for moss species richness did
not find any evidence for such a pattern (Mutke &
Barthlott, 2005; Mutke & Geffert, 2010; Shaw et al.,
2005, 2011; Tan & Pocs, 2000). We conclude that it is
too early to draw any final conclusions in this matter.
The dataset presented here offers a starting point to
further assess this question. However, what is
undoubtedly needed is more bryological field work
and collection in tropical regions of the world.

Implications for conservation

National assessments and Red Lists in industrialized
countries have shown that the rate of confirmed
extinction of bryophytes, in most cases, ranges from 2
to 4%, and that a substantial proportion of the
bryoflora worldwide is threatened in the short term
(Hallingback & Tan, 2010). Lack of resources and
funding, mean that information like this is limited to
few countries.
With limited data on the detailed local distribution
of individual species, especially in the tropical zone,
an identification of priority areas for bryophyte
diversity seems to be the most feasible approach.
Our results indicate that the centres of moss species
diversity (northern Andes, Southeast Asia, Mexico,
and Japan, as well as, the Himalayan region,
Madagascar, the East African Highlands, central
Europe, Scandinavia, and British Columbia) do not
necessarily coincide with centres of high overall
biodiversity. These places require special attention
through local assessments regarding their rare species, threat status, and possible conservation actions.
Davis et al. (1990) stated that Because many
experts believe that the threat to biological diversity
has reached a crisis stage it is our opinion that it is
unrealistic to postpone action on preserving biodiversity until complete information is collected.
Rather we must make effective use of what we
already know, while systematically organising and
expanding our knowledge base. This statement
seems to be particularly fitting for bryophytes.

Acknowledgements
An earlier version of this dataset had been established
with the help of Holger Kreft. Boon Chuan Ho,
Daud Rafiqpoor, Dietmar Quandt, and Eberhard
Fischer provided valuable assistance in obtaining
checklist publications. We thank these colleagues and
three anonymous reviewers as well for valuable
comments on this study. Parts of our work were

Global moss diversity

supported by the Akademie der Wissenschaften und

Literatur zu Mainz in the context of the long term
project Biodiversity in change (W. Barthlott) and by
the Wilhelm Lauer Foundation.
Taxonomic Additions and Changes: Nil.

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