Biol Invasions (2013) 15:1933–1946

DOI 10.1007/s10530-013-0422-2

ORIGINAL PAPER

Telling a different story: a global assessment of bryophyte

invasions
Franz Essl • Klaus Steinbauer • Stefan Dullinger •

Thomas Mang • Dietmar Moser

Received: 21 September 2012 / Accepted: 30 January 2013 / Published online: 10 February 2013
Ó Springer Science+Business Media Dordrecht 2013

Abstract We assess and review spatio-temporal increased since then. Accidental import as hitch-hiker
patterns, habitat affiliations, pathways, impacts, and (34 species) or with ornamental plants (27 species)
management experience of bryophyte invasions in constitute the most important introduction pathways.
extra-tropical countries and regions (n = 82) from five We found a remarkably high contribution from distant
continents and maritime islands spanning both hemi- donor regions to alien bryophyte floras, especially from
spheres. Distribution data were extracted and critically the complementary hemisphere. Most alien bryophytes
checked from a wide range of sources and supplemented prefer strongly modified habitats (e.g. ruderal vegeta-
with data on biology and introduction history. We tion, roadsides, lawns), and only few natural ecosystems
identified 139 bryophytes species which we consider to (forests, rocks) are regularly invaded. Evidence for an
be alien in at least one of our study regions (106 mosses, ecological impact of bryophyte invasions is scarce and
28 hepatics and 5 hornworts). Numbers of average alien competitive replacement of native moss species, or
bryophyte species are significantly higher on islands vascular plant seedlings, by alien bryophytes has only
than in continental regions of similar size, and peak been documented in a few cases. We conclude that
on maritime islands. Cumulative numbers of first bryophytes differ profoundly in many respects from
records have grown slowly until 1950 and have strongly vascular plants, and so do their invasion patterns at large
scale. Our global bryophyte invasion state assessment
provides the basis for future, more explicit consider-
Electronic supplementary material The online version of
ations of this largely neglected taxonomic group in
this article (doi:10.1007/s10530-013-0422-2) contains invasion ecology, a step we suggest to be urgently
supplementary material, which is available to authorized users. needed as studying them might provide novel insights
into patterns and processes of plant invasions in general.
F. Essl (&)
K. Steinbauer
D. Moser
Environment Agency Austria, Spittelauer Lände 5,
1090 Vienna, Austria Keywords Alien species
Bryophyta
Ecosystem

e-mail: franz.essl@umweltbundesamt.at Hemisphere
Introduction
Invasion
Naturalization

Pathway
Region of origin
S. Dullinger
T. Mang
Department of Conservation Biology, Vegetation
and Landscape Ecology, Faculty Centre of Biodiversity,
University of Vienna, Rennweg 14, 1030 Vienna, Austria Introduction
S. Dullinger
T. Mang
D. Moser
Vienna Institute for Nature Conservation and Analyses, Human activities are currently changing the distribu-
Giessergasse 6/7, 1090 Vienna, Austria tion of biota at an unprecedented scale (Ellis et al.

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1934 F. Essl et al.

2012). In particular, an ever increasing number of covering a wide range of climatic and other environ-
alien species is expanding their ranges into areas mental factors, and from different biogeographic
where they were not naturally occurring (Pyšek et al. settings (i.e. continents). As the knowledge of bryo-
2010; Essl et al. 2011) impacting native biodiversity as phyte distributions (Mutke and Geffert 2010) and
well as human welfare, and economy (Vilà et al. invasions (Essl et al. 2011) is skewed towards extra-
2011). Consequently, research in invasion biology has tropical regions with a strong tradition in floristic and
intensified during the last decades, but major taxo- taxonomic research, we excluded tropical continental
nomic and geographic gaps still remain, thus inhibit- regions. However, a few subtropical and tropical
ing a balanced understanding of mechanisms and islands (Hawaii, Ascension, St. Helena) have been
patterns driving invasions (Pyšek et al. 2008). included, as their alien bryophyte flora is well
Bryophytes (Bryophyta) include mosses (Bryopsida), studied. From the list of remaining candidate
liverworts (Hepaticopsida) and the species poor horn- regions we retained those for which comprehensive
worts (Anthoceratopsida) (Söderström et al. 2002; and up-to-date data sets (see below) on the distribution
Matteri 2003; Hill et al. 2006). In contrast to vascular of bryophytes were available. In total, we selected 82
plants, for which invasion patterns and driving forces are temperate and subtropical regions (Fig. 1, Appendix
extensively studied, global bryophyte invasions have so S1). Fifty-seven of these regions were located in the
far attracted little attention (Essl and Lambdon 2009; Northern Hemisphere (European countries and
Miller 2009; Essl et al. 2011). Generic findings for islands, US federal states, East Asian nations) and 25
vascular plants invasions should not be uncritically in the Southern Hemisphere [provinces, federal states
transferred to bryophytes because these two taxonomic or islands of South Africa, southern South America,
groups differ strongly in essential traits relevant to Oceania (incl. New Zealand and Australia), and
invasiveness: (1) bryophytes are dispersed efficiently by maritime islands]. For large countries (e.g. USA, Aus-
spores and have hence much greater natural dispersal tralia, New Zealand), we used sub-national regions in
ability than most vascular plants; (2) they have little order to refine spatial resolution in our data set.
socio-economic importance and are therefore rarely Study area size varies roughly between *5,000 and
introduced on purpose; (3) their possible effects on 250,000 km2, although some islands were signifi-
native biodiversity as well as on ecosystem functions cantly smaller (Appendix S1). For each region, we
and services are hardly known. Hence, bryophytes further collected data on region area (km2), human
invasions represent a largely unexplored chapter of population density (in the year 2000, taken from
invasion biology, and studying them can provide novel CIA 2009), and each region was classified as either
insights into patterns and processes of plant invasions. continental or as an island.
We aim for the first global assessment of bryophyte
invasions by using data from 82 regions from five Species data and analyses
continents and maritime islands on both hemispheres.
Specifically, we ask for: (1) the major taxonomic and The taxonomic concept was taken from the treatment
geographic patterns as well as temporal trends of in the most authoritative continental floras: nomen-
bryophyte invasions, (2) the prime regions of origin of clature and taxonomy follow the Bryophyte Flora of
invasive bryophytes and dominant introduction path- North America (BFNA Editorial Committee 2011) for
ways, (3) the preferably invaded habitats, and (4) the North American taxa, the most recent checklists of
main impacts, as well as (5) the available management European liverworts (Söderström et al. 2002) and
experiences and options. mosses (Hill et al. 2006) for European taxa, and
standard floras (e.g. Noguchi 1987–1994; Beever
et al. 1992; Magill and van Rooy 1998; Australian
Methods Biological Resource Study 2006) for the taxa of all
other regions. We included only accepted species into
Study regions the analysis (using the PlantList database, http://www.
theplantlist.org) because of inconsistent taxonomic
We selected a set of study areas in a two-step treatment and poor data coverage at the infra-specific
approach: first, we compiled a list of potential regions level.

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A global assessment of bryophyte invasions 1935

Fig. 1 Locations of the 82 study regions (shaded), the total number of alien and cryptogenic (a) and of naturalized (b) bryophyte
species recorded in each of them

We compiled a database on the distribution of alien bryophyte invasions (e.g. Klinck 2009; Miller 2009;
bryophytes in our study regions (Appendix S2). We Seppelt and Cave 2011; Seppelt et al. 2011), large
supplemented this data set with further information on authoritative databases (e.g. DAISIE 2012), and standard
biology, introduction history (year of first record), floras (e.g. Noguchi 1987–1994; BFNA Editorial
habitat preference, impacts, management, and pathways Committee 2011). These data sets were supplemented
from a variety of data sources (see Appendix S3): using by an exhaustive literature search (see Appendix S3)
our database on alien bryophytes in Europe (Essl and both on the ISI Web of Science and, as far as possible, in
Lambdon 2009) we searched further key literature on non-indexed journals, as many floristic records are

123
1936 F. Essl et al.

published in the latter. Finally, we contacted ca. 25 to values expected by random chance by deriving the
regional experts (see ‘‘Acknowledgements’’) who null-distribution (representing the null-hypothesis of
checked the data set and provided additional no spatial patterns between native and alien occur-
information. rences across hemispheres) by 10,000 permutation
Based on the terminology proposed by Pyšek et al. runs where in each we randomly permuted, species-
(2004), alien bryophytes were classified for each wise, the alien data entries across all regions without
region according to their invasion status as casuals native occurrence and recalculated our test statistic
(only small, non self-sustaining populations), or count.
naturalized (at least one persisting population of Statistical analyses were carried out in R (R Devel-
considerable size). This assessment was undertaken opment Core Team 2012).
by the first author based on information provided in the
data sources. We included bryophyte species as
‘‘cryptogenic’’ (=probable alien species), when strong Results
indications for an alien status in a study region (based
on factors such as anomalous geographical distribu- Alien bryophyte species
tion, association with some means of human transport,
cf. Söderström 1992) are available, but when definite We identified 139 bryophytes species which we
evidence is lacking. In doubtful cases we followed a consider to be alien in at least one study region. These
conservative approach and chose the lowest invasion comprise 106 alien mosses, 28 hepatics, and 5 horn-
category. Species which are discussed to be merely worts. This figure also includes 41 cryptogenic species
possibly alien in a given region (e.g. Riccia crystal- (37 mosses, one liverwort, three hornworts) which are
lina, Sphaerocarpos michelii, S. texanus, Tortula strong candidates to be alien, yet evidence for certainty
freibergii in Europe, Rumsey 1992; Hill et al. 2005; is lacking.
Aulacomnium palustre in New Zealand, A. Fife, Only 18 species have been recorded as alien or
personal communication) have been excluded, as probably alien from at least five regions (Fig. 2, Appen-
were records restricted to greenhouses (mostly tropical dix S2), and only few of these (especially Campylopus
and subtropical species introduced to temperate introflexus, Kindbergia praelonga, Lunularia cruciata,
regions of North America and Europe, e.g. Marchan- Orthodontum lineare, Pseudoscleropodium purum) are
tia planiloba, Splachnobryum obtusum, Vesicularia widespread.
reticulate, Zoopsis liukiuensis; Arts 2001; Porley and
Hodgetts 2005). Biogeographic pattern and spatial distribution
Our final database included all bryophyte species
which were recorded as alien or cryptogenic in at least The numbers of alien bryophytes are rather low in
one of the 82 study regions. To test if recipient region most regions. There are only 10 (12 %) regions where
size or human pressure (population density) influence C10 alien bryophyte species were recorded (Appendix
bryophyte invasions, we related a regions’ total alien S4). These 10 regions are scattered across continents,
bryophyte species numbers with the (log-transformed) with no major hotspots of bryophyte invasion emerg-
study region area and human population density by ing so far (Fig. 1). In Europe the countries with the
means of Poisson family generalized linear models highest numbers of alien bryophytes are the UK (22
(GLMs) (Faraway 2006). To test for possible biogeo- species), followed by Ireland (11 species), and France
graphical differences, this analysis was performed (10 species). In North America, states in the pacific
separately for islands and continental regions. West (British Columbia: 13 species; California: 10
We further tested if bryophytes preferentially species), and to a lesser degree in the Northeast
invade regions of the hemisphere they are not natively (Massachusetts: 6 species) are most invaded. How-
found on through a non-parametric permutation test: ever, on the islands of Hawaii 22 alien bryophyte
as test statistic we counted all data entries of a species’ species are known. Globally, highest alien bryophyte
alien occurrence in regions of one hemisphere while numbers were recorded on the islands of the Southern
species’ native regions are yet entirely restricted to the Hemisphere, with the South and North Island of New
complementary hemisphere. We compared this count Zealand (27 species each) and St. Helena (22 species)

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A global assessment of bryophyte invasions 1937

dated records are available and hence no trend can be

derived).
No alien bryophyte species is known of having
arrived in any of our study regions prior to 1800, but
this is unsurprising as bryology as discipline evolved
relatively late (and few common species had received
their modern scientific names by this date). Most early
records stem from Europe; the first species to be
recognized as introduced were L. cruciata (1828 in
Karlsruhe, Germany; Frahm 1973) and Atrichum
crispum (1848 in Rochdale, England; Smith 2004).
The first records outside Europe were made in the late
nineteenth century: Phaeoceros carolinianus and
Fig. 2 The 18 alien bryophyte species which have been P. purum both in 1875 on St. Helena (Dickson
recorded in at least 5 study regions (out of 82). Different 1967), and Pleuridium subulatum var. subulatum in
invasion status is shown in black (naturalized), dark grey 1879 on the South Island of New Zealand (A. Fife,
(casual), and cryptogenic (light grey) personal communication).
Due to its rapid speed the colonization of Europe by
being most invaded. The continental regions of South
C. introflexus represents one of the best documented of
America, Asia and Africa have much lower numbers
all plant invasions (Hassel and Söderström 2005;
of alien bryophytes: four species (regions VIII and IX
Klinck 2009; Mikulášková et al. 2012). It was first
in Chile) in South America, six species (Japan) in East
recorded in England in 1941, and by 1950 already
Asia, and three species (Western Cape Province, South
approximately 60 invaded sites were known in the UK.
Africa) in Africa. In four mid- and south-western
The first record of this species on continental Europe
states of the USA no alien bryophytes were recorded,
was made in 1954 on the channel coast of Bretagne
and in 27 regions only one or two alien bryophytes are
(Finistère) (Richards 1963; Hassel and Söderström
known.
2005); within a year it was then found near Paris
Despite a globally scattered distribution of alien
(Fontainebleau), and subsequently it spread rapidly to
bryophyte records, average alien bryophyte species
neighbouring countries. Today its known distribution
numbers vary substantially across continents, being
stretches to Russia in the east, to the fringes of the
highest in Oceania (Table 1). Moreover, numbers of
Mediterranean basin in the south and also includes the
alien bryophyte species on maritime islands are much
Azores (Klinck 2009). Other well-documented exam-
higher than in continental regions of similar size
ples of fast spread include Orthodontium lineare in
(Fig. 3): while region area significantly influences
Europe (Hassel and Söderström 2005), Ptychomitrium
total alien bryophyte species numbers for both conti-
serratum (Miller and Robinson 2010) and P. purum in
nental regions as well as maritime islands (islands:
North America (Miller and Trigoboff 2001; Miller
p \ 0.001; continents: p \ 0.001), no such pattern
2009), and K. praelonga and P. purum on Tasmania
was found for human population density (Fig. 3)
(Seppelt and Cave 2011; Seppelt et al. 2011).
(islands: p = 0.14; continents: p = 0.10).
Origins of alien bryophytes
Temporal trends
The majority of alien bryophytes have large native
For 293 records (71 %) information on a species’ first ranges encompassing several continents, with 75
record in a region was available. The analysis of the species (54 %) native to at least three continents. In
temporal invasion patterns shows that numbers of first order of decreasing importance, most species which
records have grown slowly until 1950, but since then have become alien are native to North America (86
have increased strongly without any sign of saturation species), followed by Europe (78 species), South
so far (Fig. 4). This is true for both the global level as America (62 species), temperate Asia (57 species),
well as all individual continents (for Africa only two Africa (50 species), and Australasia (41 species). For

123
1938 F. Essl et al.

Table 1 Means and standard deviations (in brackets) of bryophyte species numbers across regions of different continents (numbers
of regions per continent in parentheses)
Cryptogenic Casual Naturalised Total

Europe (n = 35) 2.0 [1.4] 1.0 [1.0] 2.5 [3.5] 5.5 [4.6]
North America (n = 19) 0.7 [1.8] 1.8 [1.4] 1.2 [1.3] 3.6 [3.4]
Asia (n = 3) 0.7 [1.2] 0.7 [1.2] 1.0 [1.0] 2.3 [3.2]
Oceania (n = 8) 2.4 [1.8] 3.4 [2.8] 6.3 [7.9] 12.0 [10.0]
South America (n = 9) 0.0 [0.0] 1.1 [0.6] 1.2 [0.8] 2.3 [1.1]
Maritime islands of S. Hemisphere (n = 5) 0.4 [0.9] 2.0 [3.4] 3.2 [6.1] 5.6 [9.2]
Africa (n = 2) 0.0 [0.0] 1.5 [0.7] 0.5 [0.7] 2.0 [1.4]
Values are provided both for total numbers as well as separately for subsets pertaining to different invasion status categories. Hawaii
has been excluded from the analysis

Fig. 3 Total alien bryophyte species numbers per region in whether the region is an island or not (GLM; for human
relation to expected values depending on region area population density the value was fixed to the mean). Grey dots
(a) (islands: p \ 0.001; continents: p \ 0.001), human popula- denote islands, black circles continental regions
tion density (b) (islands: p = 0.14; continents: p = 0.10), and

eight species the region of origin is unknown or

controversial (Fig. 5).
Bryophytes which are exclusively native to one
hemisphere significantly more frequently invade
regions on continents of the complementary (p =
0.02) (Table 2). For example, C. introflexus and
O. lineare, which are by now widely distributed in
the Northern Hemisphere, are both of southern hemi-
spheric origin. On the contrary, in North America
some of the more widespread bryophytes are Euro-
pean species (e.g. Brachythecium albicans, L. cruci-
ata, P. purum, Thuidium tamariscinum). In temperate
regions of the Southern Hemisphere, naturalized
introduced bryophytes include the pleurocarp Euro-
Fig. 4 Invasion curve of alien bryophytes given as cumulative pean mosses P. purum (Tasmania, New Zealand,
numbers of first records in the 82 study regions. Given is the
South America, St. Helena), R. squarrosus (Tasmania,
invasion curve for each continent (Africa being excluded as only
two records were available). For numbers of study region per New Zealand), R. triquetrus and Sphagnum subnitens
continent see Appendix S1 (the latter two in New Zealand).

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A global assessment of bryophyte invasions 1939

2009) and northern Italy (Hypopterygium tamarisci;

Aleffi et al. 2010), or tropical plants to a nursery in
northern Italy (Sematophyllum adnatum; Frey et al.
2006). Some literature sources identify certain species
as being introduced ‘‘with soil’’ (Paton 1999), but this
may often also refer to soil in plant pots. It has been
argued that the UK record of Lophozia herzogiana in
Woolmer Forest can be attributed to unintentional
introduction with imported wool from New Zealand
(Australian National Botanic Garden 2012). Only four
species are known to having been introduced deliber-
ately (e.g. Ricciocarpos natans, which is floating in
water and hence used for garden ponds and aquaria, in
Fig. 5 Observed (dashed line) total count of all alien occur- northern Europe and Hawaii). In western North
rences on a given hemisphere if species’ native distributions are America and elsewhere (e.g. St. Helena; Dickson
exclusively restricted to the complementary hemisphere, 1967), P. purum has been introduced intentionally in
compared against the null-distribution derived by permutation
of alien entries. Observed bryophyte invasions occur more often packing material for nursery stock (Schofield 1997),
on the complementary hemisphere than what can be expected by and it is likely to be occasionally transported as a
random chance (p = 0.02) contaminant of peat-based compost. However, for
51 % of the alien bryophytes their mode of introduc-
Pathways tion remains unknown.
Secondary spread mediated by wind-dispersal is
The most important introduction pathway of alien assumed to be decisive for range expansions over
bryophytes is unspecified accidental import as a hitch- large distances in some cases (e.g. the spread of
hiker on ships and planes, and perhaps on clothing or C. introflexus in Europe, Klinck 2009; or the recent
goods (34 species). The only other significant intro- spread of P. serratum to the northeastern USA, Miller
duction pathway is as an epiphyte on ornamental and Robinson 2010). In few cases, recent introductions
plants or as a weed in the transport of compost and of bryophytes associated with wetlands are sus-
horticultural supplies (27 species), e.g. in association pected to having occurred through migratory birds,
with the import of Australasian tree ferns to Ireland e.g. Riccia fluitans in Hawaii (Hall et al. 1995) and
(Calomnion complanatum, Dicranoloma menziesii, Micromitrium megalosporum in Japan (H. Deguchi,
Leptotheca gaudichaudii; Holyoak and Lockhart personal communication).

Table 2 Regions of origin of the alien bryophyte species recorded in the 82 study regions on the different continents
Continents (# regions) # species Europe Temp. Asia Trop. Africa Australasia N. America S. America Unknown
Asia

Europe (n = 35) 48 10 11 6 14 19 20 23 4
North America (n = 19) 23 15 12 7 11 9 12 13 0
Asia (n = 3) 27 14 11 8 11 8 21 14 2
Oceania (n = 8) 46 44 30 22 22 14 36 19 0
South America (n = 9) 8 8 6 4 5 3 6 4 0
Maritime islands of 25 18 13 13 12 17 17 14 2
S. Hemisphere (n = 5)
Africa (n = 2) 3 3 1 0 1 1 1 1 0
Given are the total number of alien bryophyte species recorded on a continent (# alien bryophyte species), and species’ native ranges
have been assigned to continents. Species which inhabit several continents have been assigned to multiple regions of origin. Hawaii
has been excluded from the analysis

123
1940 F. Essl et al.

Invaded ecosystems wood in woodlands (Hedenäs et al. 1989; Herben

1994; Hassel and Söderström 2005), and aside from
Across all continents, alien bryophyte species have a anthropogenic habitats L. cruciata invades moist,
clear affinity to habitats with a strong anthropogenic shaded soils and rocks. In Australia and New Zealand
disturbance regime (Table 3). Most frequently colo- some northern hemispheric species (e.g. K. praelonga,
nized are gardens (41 species), followed by road- and P. purum, R. squarrosus, R. triquetrus; Seppelt and
tracksides (34 species), other ruderal habitats (23 Cave 2011; Seppelt et al. 2011; A. Fife, personal
species), and broadleaved forests (21 species). In communication) have locally invaded native bushland
particular in North America, Tasmania and New and forest vegetation, whereas S. subnitens is invading
Zealand, several bryophyte species frequently colo- wetlands on the west coast of New Zealand (Dobson
nize lawns and grasslands (20 species, e.g. Brachy- 1975; Landcare Research 2012). Few bryophytes (e.g.
thecium spp., P. purum). The selection of substrates Fontinalis taxifolius, F. antipyretica in South Africa,
underlines the preference of disturbed habitats. R. fluitans in Europe, R. natans in Europe and Hawaii;
Exposed soil, which is created by mechanical distur- Miller 1967; Hall et al. 1995; Magill and van Rooy
bance, is most often recorded (81 species), followed 1998) have locally invaded aquatic ecosystems.
by rocky substrates (23 species); other micro-habitats
are only rarely colonized.
Few bryophytes have so far widely established in Ecological and economic impacts
natural, or near-natural vegetation (Söderström 1992;
Stieperaere and Jacques 1995). In Europe and western The impacts of alien bryophytes are less obvious than
North America C. introflexus colonizes mainly coastal those of alien vascular plants. Their small size makes
dunes, heathlands and bogs, but recently also conifer them weak competitors for light, nutrients or space,
plantations (Mikulášková et al. 2012). Its occurrence except of native bryophytes, lichens or seedlings of
in the former, near-natural habitats however is often vascular plants; only rarely have been effects on
associated with burned areas, peat cuttings or roadside animal species, ecosystem structure and functioning
banks. Also in Europe O. lineare occurs on decaying observed.

Table 3 Habitat affiliation by alien bryophytes in the 82 study regions

Habitat groups Habitats invaded No. %

Heavily modified vegetation Fields 12 4.5

Heavily modified vegetation Ruderal vegetation 23 8.7
Heavily modified vegetation Track- and roadsides 34 12.9
Heavily modified vegetation Greenhouses 10 3.8
Grasslands Grasslands, lawns 20 7.6
Grasslands Gardens, parks, cemeteries 41 15.5
Grasslands Dunes 3 1.1
Rocks and walls Rocks 13 4.9
Rocks and walls Walls 14 5.3
Forests Conifer forest 8 3.0
Forests Broadleaved forest 21 8.0
Wetlands Mires, bogs 8 3.0
Wetlands Pioneer vegetation of water bodies 19 7.2
Freshwater Standing freshwaters 3 1.1
Unknown Unknown 35 13.3
Total 264 100
Given are numbers of species per habitat and the share of the total number of habitat records. Note that in many cases species occur in
more than a single habitat. For 35 species their habitat affiliation is unknown

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A global assessment of bryophyte invasions 1941

The impacts of C. introflexus in Europe and thriving ‘‘moss-killer’’ industry based on herbicide
P. purum in North America and the Southern Hemi- application (Schofield 1997).
sphere are the best studied. Both species are capable of In a few cases, precautionary management has
forming dense mats. C. introflexus significantly aimed at eradicating newly introduced bryophytes to
reduces species diversity of other lower plants and limit future impacts. On the South Island of New
lichens in dunes and nutrient-poor grasslands (Bier- Zealand efforts are underway to eradicate recently
mann and Daniels 1997; Ketner-Oostra and Sýkora introduced Rhytidiadelphus triquetrus (Landcare
2004, 2008; Hasse 2007), reduces the recruitment of Research 2012), as is the case for Fissidens taxifolius
Calluna vulgaris (Equiha and Usher 1993), and on the Chatham Islands (A. Fife, personal commu-
negatively affects species diversity of grasshoppers nication).
(Schirmel 2011), soil-entomofauna (Vogels et al.
2005), carabids and spiders in dunes of the Baltic
Sea (Schirmel et al. 2011). P. purum is considered to Discussion
change species composition in invaded ecosystems,
e.g. in bushland of higher elevations on St. Helena Alien bryophytes: species diversity, numbers,
(P. Lambdon, personal communication), Hawaii and spatial patterns
(Volcanoes National Park, Waite 2007), and Tasmania
(Seppelt et al. 2011). A few other northern hemi- This first global assessment on bryophyte invasions
spheric pleurocarp mosses (K. praelonga on St. reveals that the number of bryophyte species which
Helena, R. squarrosus on Tasmania, Rhytidiadelphus have become alien somewhere in the 82 study regions
triquetrus in New Zealand; A. Fife, P. Lambdon and is moderate. Given the broad geographic coverage of
R. Seppelt, personal communication) are considered the study regions (with the notable exception of
to outcompete native bryophytes in selected regions of tropical regions), this pattern is very likely to be valid
the Southern Hemisphere, however studies on the globally.
magnitude of this impact are lacking. Of the 15,344 bryophyte species which are in
A few threats posed by other alien bryophytes have current use or without recent synonymy in the
been documented, but so far the implications have not PlantList database (the most comprehensive standard
been well studied yet. In the UK, O. lineare competes source on taxonomy and nomenclature of bryophytes),
with a rare relative, O. gracile, on sandstone rocks and we found 137 being alien or probably alien (crypto-
in many cases has caused the local loss of this species genic) (0.9 % of all species). In Europe, the continent
(Porley and Hodgetts 2005). Similarly, Lophocolea with the best studied native and alien bryophyte flora,
semiteres may be ousting the native L. heterophylla in this proportion is similar: Hill et al. (2006) record
the Netherlands (Porley and Hodgetts 2005; Porley 1,292 species of mosses and Söderström et al. (2002)
and Haynes 2009). record 474 species of liverworts (excluding subspecies
and varieties) as native. On this basis, only 1.8 % of all
European species are certainly alien; if cryptogenic
Management experience and options species are included, the estimate rises to 2.5 %. Of all
study regions, the most striking exception to this
Little experience on managing bryophyte invasions is pattern is St. Helena, where 22 alien bryophyte species
available so far. Tested management strategies include constitute approximately one-fifth of the total native
liming of dunes, burning, herbicide treatment and bryoflora (P. Lambdon, personal communication).
introduction of grazing animals to trample the mats of The low number of alien bryophytes strongly
C. introflexus in western European dunes (Klinck contrasts with the much greater proportion of aliens
2009). These efforts have generally yielded only among vascular plants (e.g. Weber 2003; Lambdon
limited success, and in most cases such direct control et al. 2008, Pyšek et al. 2008). One reason for the
measures risk damaging native vegetation more than relative paucity of bryophyte invasions is that due to
they affect the invader (Ketner-Oostra and Sýkora the lack of (historical) distribution data (Magill 2010;
2000; Klinck 2009). The invasion of P. purum in urban Mutke and Geffert 2010), some alien bryophytes
lawns in the western and eastern USA has led to a (especially inconspicuous species) might have been

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overlooked and may thus wrongly be considered the islands habitats (e.g. Denslow et al. 2009; Kueffer
indigenous. This especially applies to earlier intro- et al. 2010); for bryophytes however we additionally
ductions, where the status is sometimes difficult to suggest island systems to be particularly suited for
determine. Further, Mutke and Geffert (2010) have invasions as climates tend to be more humid, mois-
recently shown that European countries and subna- ture-saturated and less exposed to extreme climatic
tional administrative units have, on average, recorded amplitudes.
significantly higher numbers of bryophytes than extra- Although the tropics have not been covered in this
European counterparts, which is an indication that the assessment due to paucity of data, some bryophyte
bryophyte flora of many extra-European regions may species are assumed to be alien in tropical regions, and
be considerably under-documented. in particular in man-made habitats in tropical moun-
More important than possible data biases however, tain ranges (Gradstein and Vana 1987; Norris et al.
spores of bryophytes are very efficient at long distance 1999). However, it has to be left to future research
dispersal (Porley and Hodgetts 2005). This implies that based on an improved data basis to assess the
we must expect human activities to play role of much importance of bryophyte invasions in tropical regions.
lesser importance in overcoming geographic barriers
than in vascular plants. Partly for this reason, bioge- Regions of origin
ographically anomalous distributions are less reliable
as indicators of human introductions than in less Compared to other taxonomic groups and in particular
mobile taxa. The occurrence of Scopelophila catarac- vascular plants, the important contribution of distant
tae in Europe, which is confined to mine spoil heaps donor regions (especially from the other hemisphere)
contaminated by heavy metals, is a famous example: it to a continents’ alien bryophyte flora is remarkable. In
was first recorded in 1967 in Wales and has few known Europe, 65 % of the 48 alien bryophyte species are
European localities which are all very distant from native to the Southern Hemisphere. In contrast, just
each other (Smith 2004; Colpa and van Zanten 2009). 19 % of alien bryophyte species are native to other
Its main range is in the Southern Hemisphere, and it is parts of this continent, whereas for vascular plants the
difficult to imagine how it reached these European figure is 53 % (Lambdon et al. 2008). In North
outposts, whether by human or natural means. We America, again 65 % of the 23 alien bryophytes are
include this species as cryptogenic. native to the Southern Hemisphere, while 52 % are
Region area is a significant predictor of alien native to this continent. For Oceania, 93 % are native
bryophyte species numbers, but interestingly the to the Northern Hemisphere, whereas only 30 % of the
number of alien bryophyte species in a study region 44 species are native to this region.
shows no significant relationship with human pressure We argue that the strong contribution of alien
(population density). Instead, other factors seem to be bryophyte species from the other hemisphere in conti-
more relevant in explaining geographic patterns of nental alien floras is linked to the excellent dispersal
bryophyte invasions: first, Essl and Lambdon (2009) capacities of bryophytes, which makes it easier to
found that European regions with a humid climate are overcome geographical barriers within a hemisphere
most invaded, a pattern which might hold true for our without the assistance of man (Porley and Hodgetts
study regions. Such a pattern is also reflected in native 2005). However, crossing the equator by natural means
bryophyte species numbers in a similar way (Mutke presents a barrier because of the prevailing wind
and Geffert 2010), suggesting that climatic conditions directions in the intertropical convergence zone (Barry
impose a strong filter on bryophyte invasions. Second, and Chorley 1992). Only through the activities of man,
numbers of alien bryophytes on islands are on average in particular unintentional human-aided transport of
significantly higher than numbers on continental spores or live bryophytes, this distribution barrier can
regions of similar size. Higher alien species numbers now be overcome much easier.
on islands are known to be a consistent feature for
most, if not all major plant and animal taxonomic Pathways
groups (e.g. Cronk and Fuller 1995; Reaser et al. 2007;
Denslow et al. 2009), with differing levels of invasion The low importance of deliberate introductions
being closely related to the extent of human impact on strongly contrasts with what is known from vascular

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A global assessment of bryophyte invasions 1943

plants, where intentional introduction contributes most strongholds in anthropogenic habitats (Essl and
to alien species numbers (Hulme et al. 2008). Indeed, in Lambdon 2009); on the other hand, 27 % of the alien
Europe 63 % of all established alien vascular plant bryophyte species are entirely constricted to and
species have been introduced intentionally (Lambdon another 36 % are predominantly occurring in anthro-
et al. 2008), whereas only one alien bryophyte species pogenic habitats.
has been introduced to this continent on purpose. Generally, the habitat affiliation patterns resemble
The velocity of secondary spread within the alien that of vascular plants (e.g. 64 % of established alien
range often mainly depends on the dispersal capacity of vascular plants in Europe invade ruderal habitats;
the species, but human activity may enhance it further. Lambdon et al. 2008), but alien bryophytes are even
Spread may be promoted by anthropogenic changes to more restricted to man-made habitats, as in vascular
existing habitats (e.g. by airborne pollutants), by plants grasslands and woodlands (and here in particular
creating new habitats, or unintentional transport. Some riverine forests) (37 % respectively 32 % of all estab-
alien bryophytes are suspected to have expanded in lished alien vascular plants in Europe; Lambdon et al.
West and Central Europe due to changed chemical 2008) are also heavily invaded (cf. Chytrý et al. 2005).
characteristics of substrates due to acid rain and airborne
nitrogen deposition (e.g. C. introflexus, O. lineare; Future expected trends
Söderström 1992; J.-P. Frahm, personal communica-
tion). Ketner-Oostra and Sýkora (2000) have shown that Man-made modifications of existing and creation of
the spread of C. introflexus in Europe is fostered by new habitats, and increasing intercontinental trade
mechanical disturbance of invaded habitats (e.g. caused are the dominant drivers for the spread of alien
by rabbits or trampling by people), which breaks up the bryophytes. Furthermore, climate change and rising
moss mat and subsequently disperses propagules to temperatures may foster future range expansions of
initiate new populations. The formation of specialized alien bryophytes (Frahm and Klaus 2001). In Europe,
structures for vegetative reproduction is a potential some (sub-)tropical bryophyte species formerly
mechanism to aid invasion via vegetative fragments, restricted to greenhouses in Europe have recently
gemmae (simple asexual buds shed from the leaf tips or started to also becoming established outdoors
other organs), bulbils, or tubers (larger gemmae-like (e.g. Didymodon australasiae; Kucera 1999; Müller
structures borne respectively in the leaf axils or on 2002), which might reflect the trend of global
rhizoids). More than 50 % of UK alien bryophytes atmospheric warming.
possess these structures, compared with only 28 % of As inter-continental trade increases further, and
natives (Essl and Lambdon 2009). However, many especially with the rising popularity of exotic orna-
bryophytes are dioecious, and as there is often only one mental plants (Pemberton and Liu 2009), there is a
sex in the founder population (e.g. Leptophascum great potential for increased movement of alien
leptophyllum, L. cruciata in their alien European range), bryophyte species across the world. While airborne
their spread is restricted to vegetative propagation. acidification has been strongly reduced in the last few
decades and it is hence likely that this factor will lose
Habitat preferences relevance, nitrogen deposition is still increasing and
exceeds critical loads in industrialized regions of
Most alien bryophytes are early-successional colonists Europe and North America (Bobbink et al. 2010), and
of strongly modified habitats (e.g. ruderal vegetation, as several alien bryophyte species can take advantage
roadsides, lawns) while natural ecosystems are rarely of airborne eutrophication (Söderström 1992) they
invaded, and a range of natural ecosystems in which will clearly benefit from this trend.
bryophytes comprise essential components are very Deriving feasible management strategies for bryo-
rarely (e.g. mires) or not at all (e.g. alpine vegetation, phytes to control further spread is very difficult, for
screes) invaded. This is in strong contrast to the habitat several reasons. Introduction is difficult to control,
preferences of native bryophytes: for example, of identification of species needs expert knowledge,
1,027 species recorded as native in the UK, only 89 long-distance dispersal and thus re-immigration is
species (9 %) are considered to be strongly adapted to likely to be frequent, and due to the small size of
agricultural habitats, and only 31 (3 %) have their core bryophytes, most management measures are difficult

123
1944 F. Essl et al.

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