Global Moss Diversity

Global moss diversity: spatial and taxonomicpatterns 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 400different geographical units and standardized species taxonomy using the TROPICOS database of theMissouri Botanical Garden. Maps of overall moss species richness, as well as individual maps for taxonomicorders of mosses, are provided. Based on our dataset, we did not find a general latitudinal gradient ofincreasing moss diversity with decreasing latitude. Several areas of temperate broadleaf forests, borealforests, and tundra show relatively high species richness that is comparable to tropical regions. Centres ofmoss diversity include the northern Andes, Southeast Asia, Mexico, and Japan, as well as the Himalayanregion, Madagascar, the East African Highlands, central Europe, Scandinavia, and British Columbia. Ourdataset presents the first collection of moss species inventories with global coverage. It contributes todocumentation and understanding of global biogeographic patterns in mosses, helps to identify gaps infloristic knowledge, and could prove to be a valuable resource to aid taxonomic and systematic revisions orassessments of species and genera, by quickly and easily supplying an overview of the geographicdistribution of a given taxon.

Keywords: Biogeography, Bryophyte diversity, Latitudinal gradient, Macroecology, Moss species richness

IntroductionSpecies diversity is unequally distributed across the

globe (Barthlott et al., 2007). Exploration and doc-

umentation 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,

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 catalo-

guing 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 catalo-

guing completeness in many tropical countries have

increased (Shaw et al., 2005), actual analyses on

continental or global scales are still sparse.

Conservation of bryophyte diversityBiodiversity is rapidly declining across the globe

(Barnosky et al., 2011). This is increasingly raising

concerns amongst scientists and policy makers, and

Correspondence to: Jan Laurens Geffert, Nees-Institut fur Biodiversitat derPflanzen, Rheinische Friedrich-Wilhelms-Universitat Bonn, MeckenheimerAllee 170, 53115 Bonn, Germany. Email: [email protected]

British Bryological Society 2013DOI 10.1179/1743282012Y.0000000038 Journal of Bryology 2013 VOL. 35 NO. 1 1

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 diversitySo 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, how-

ever, 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.

MethodsData collectionWe collected moss species inventories of political or

regional units worldwide (hereafter geographical

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 Source

Antarctica Ochyra et al., 2008Argentina Matteri, 2003Australia Klazenga et al., 2009Austria Kockinger et al., 2008Bangladesh OShea, 2003aBelgium Sotiaux et al., 2007bBhutan Long, 1994Bolivia Churchill et al., 2009Brazil Forzza et al., 2010Canada Ireland et al., 1987Central andSouth America

Delgadillo et al., 1995

Central Europe Hill et al., 2006Chile Muller, 2009China Redfearn et al., 1994Corsica Sotiaux et al., 2007aEstonia Ingerpuu et al., 1994Former Soviet Union Ignatov et al., 2006France Lemonnier, 2010Galapagos Ziemmeck and

Gradstein, 2009Germany Koperski et al., 2000Great Britain Hill et al., 2008Greenland Goldberg, 2003Guianas Boggan et al., 1997Hawaii Staples et al., 2004Hong Kong Hu et al., 2002Hungary Erzberger and Papp, 2004Iceland Johannsson, 2003India Lal, 2005Indochina Tan and Iwatsuki, 1993Japan Iwatsuki, 2004Korea Park and Choi, 2007Liechtenstein Senn, 2000Luxembourg Werner, 2008Malesia Tan and Soon, 1997Middle East Frey and Kurschner, 1991Mongolia Tsegmed, 2001Netherlands Siebel et al., 2005New Caledonia Pursell and Reese, 1982New Zealand Fife, 1995North Africa Ros et al., 1999Pakistan Higuchi and Nishimura, 2003Seychelles Frahm and Ho, 2009Slovakia Kubinska and Janovicova, 1998South Africa Germishuizen and Meyer, 2003Southeast Europe Sabovljevic et al., 2008Sri Lanka OShea, 2002Sub Saharan Africa OShea, 2003bSulawesi Gradstein et al., 2005Sumatra Ho et al., 2006Svalbard Frisvoll and Elvebakk, 1996Switzerland Nationales Inventar der

Schweizer Moosflora, 2010Taiwan Chiang et al., 2001Thailand He, 1997United States of America The New York Botanical Garden, 2003

Geffert et al. Global moss diversity

2 Journal of Bryology 2013 VOL. 35 NO. 1

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 standardizationBecause 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 com-

promising 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 (RDevelopment Core Team,

2011). The basic mechanism is to generate a specific web

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 nomen-

clatural 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 richnessFor quantitative analysis of our dataset, the standar-

dized 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 neces-

sary 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:

S1~S2A1

A2

z

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 subgroupsCollecting species lists instead of species numbers

made it possible not only to analyse moss species


Journal of Bryology 2013 VOL. 35 NO. 1 3

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 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.

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).



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.

ResultsSpecies inventories and taxonomicstandardizationInventory 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 richnessMoss 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 majorsubgroupsThe 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

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.



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).



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

Figure 4b



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 diver-

sity 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 pantro-

pical 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 ofNewZealand, 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.

DiscussionThe 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 adapt-

ability to different environmental conditions. The low

species richness in many dry-land regions, however,

demonstrates their high dependence on freely avail-

able 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

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 richnessOur 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 sig-

nificantly 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



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 conservationNational 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 spe-

cies, 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 biodi-

versity 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.

AcknowledgementsAn 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

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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Global Moss Diversity