Freshwater Biology (2006) 51, 1879–1889 doi:10.1111/j.1365-2427.2006.01624.

Relationships between distribution and abundance vary

with spatial scale and ecological group in stream
bryophytes
JANI HEINO* AND RISTO VIRTANEN†
*Research Programme for Biodiversity, Finnish Environment Institute, University of Oulu, Oulu, Finland
†
Department of Biology, Botanical Museum, University of Oulu, Oulu, Finland

SUMMARY
1. We examined whether the local abundance of stream bryophytes in a boreal drainage
basin (Koutajoki system in northeastern Finland) correlated with their: (i) regional
occupancy; (ii) provincial distribution in northwestern Europe; and (iii) global range size.
We specifically tested whether aquatic and semi-aquatic species differ in their distribu-
tion–abundance relationships. We also analysed the frequency distributions of occupancy
at two spatial scales: within the focal drainage system and across provinces of
northwestern Europe.
2. Regional occupancy and mean local abundance of stream bryophytes were positively
correlated, and the relationship was rather strong in aquatic species but very weak in semi-
aquatic species. Local abundance was related neither to provincial distribution nor global
distribution.
3. Species frequency distributions differed between regional occupancy and provincial
distribution. While most species were rare with regard to their regional occupancy within
the focal drainage system, most of the same set of species were common and occurred in
most provinces in northwestern Europe.
4. The results indicate the presence of dominants (core species) and transients/
subordinates (satellite species) among stream bryophytes, highlighting marked differen-
tiation in life-history strategies and growth form. The observed abundance–occupancy
relationships suggest that dispersal limitation and metapopulation processes may govern
the dynamics of obligatory aquatic stream bryophytes. In semi-aquatic species, however,
habitat availability may be more important in contributing to regional occupancy.

Keywords: bryophytes, distribution patterns, interspecific distribution–abundance relationship,

streams

Blackburn, 2000). Among the most studied macro-

Introduction
A major aim of macroecology is to understand the
patterns in range size, regional distribution and
abundance of organisms, as well as the processes
generating these patterns (Brown, 1995; Gaston &
Correspondence: Jani Heino, Research Programme for
Biodiversity, Finnish Environment Institute, PO Box 413,
University of Oulu, FIN-90014 Oulu, Finland.
E-mail: jani.heino@ymparisto.fi
 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd
ecological patterns is that between regional distribu-
tion and local abundance of species. A plethora of
studies have examined this relationship both empiri-
cally and experimentally, and most have reported a
strongly positive relationship between regional dis-
tribution and local abundance of species (Collins &
Glenn, 1997; Gaston & Blackburn, 2000; Guo, Brown &
Valone, 2000; Holt, Warren & Gaston, 2004). The
abundance–distribution relationship is so prevalent in
nature that it has been considered as one of the few
1879
1880 J. Heino and R. Virtanen
true generalities in ecology (Gaston, 1999). Despite
this prevalence, the processes associated with the
generation of the positive distribution–abundance
relationship are not well understood, and several
differing processes supposedly act in concert. The
multitude of processes is also suggested by the large
number of general hypotheses coined to account for
the pattern, and to date at least nine such hypotheses
have been claimed to explain the positive distribu-
tion–abundance relationship (Gaston, Blackburn &
Lawton, 1997). These hypotheses include niche
breadth, niche position, metapopulation dynamics
and range position, as well as a few potential artefacts
related to phylogenetic non-independence and sam-
pling problems (Brown, 1984; Hanski, Kouki &
Halkka, 1993; Gaston et al., 1997).
Although much has been written about the positive
distribution–abundance relationship, few studies
have examined the influence of scale and measure-
ment of distribution on the pattern (Gaston &
Lawton, 1990; Gregory & Blackburn, 1998; He &
Gaston, 2000). That is, a measure of local abundance
may be related either to regional occupancy, regional
range or even global range size. The classic studies
on the distribution–abundance relationship concen-
trated on regional occupancy and local abundance
(e.g. Hanski, 1982), mirroring the relationships
between site-specific measures of population charac-
teristics. Further studies have related a separately
derived measure of abundance to distribution across
larger regions, for example, the number of grid cells
occupied rather than the number of sites occupied
within a region (reviewed by Gaston & Blackburn,
2000). At the extreme are the studies that have
related a measure of abundance to species global
range size that can be estimated either by area extent
or division of species in groups of differing range
size class. It is likely that such widely varying
measures of distribution could influence the strength
of the relationship between distribution and abun-
dance (Thompson, Hodgson & Gaston, 1998). Similar
to the distribution–abundance relationships, one
might expect that species distribution patterns differ
with spatial scale (e.g. McGeoch & Gaston, 2002;
Bossuyt, Honnay & Hermy, 2004).
Besides being of general interest, distribution–
abundance relationships and regional distribution
patterns of species can reveal important aspects of
an organism’s life history and functional significance
 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd, Freshwater Biology, 51, 1879–1889
in an ecosystem. An important recent advance was
made by Gibson, Ely & Collins (1999) who linked
explicitly the traditional core–satellite model of
Hanski (1982) and Grime’s (1998) dominant-subordin-
ate-transient model. In this context, the distribution–
abundance relationship may provide new insights
through combining the regional dynamics of species
and functional consequences of such dynamics. Thus,
a positive distribution–abundance relationship may
be related to the idea that species are functionally
differentiated, and that there are potentially recognis-
able groups of species that represent different strat-
egies in response to ecosystem characteristics (e.g.
Muotka &Virtanen, 1995). There is still rather little
evidence about how the distribution–abundance rela-
tionship applies to different groups of organisms (e.g.
many plant groups) in different systems and what can
be actually learnt from their examination (cf. Freckl-
eton & Watkinson, 2003).
Although the relationship between distribution and
abundance has been studied in a wide variety of
contexts, some systems and organism groups have
received rather little attention from the macroecologi-
cal perspective. Foremost among these are marine
(Wieters, 2001; Foggo, Frost & Attrill, 2003; Frost et al.,
2004) and freshwater organisms (Pyron, 1999; Tales,
Keith & Oberdorff, 2004), and a comparison of the
established macroecological patterns of terrestrial
organisms with those of aquatic organisms would
certainly increase our understanding of the true
generality of the supposedly ‘general’ patterns, as
well as the proposed underlying mechanisms. One
amenable model group for examining the distribu-
tion–abundance relationship in aquatic ecosystems
are stream bryophytes (see Muotka & Virtanen, 1995).
Stream bryophytes live in spatially and temporally
heterogeneous habitats, characterised by frequent
disturbances and relatively high connectivity among
sites within a drainage system. Thus, one might
expect that sites within a drainage basin are fre-
quently denuded of and recolonised by bryophytes,
eventually leading to the positive distribution–abun-
dance relationships via metapopulation dynamics
(e.g. Hanski, 1982). Similar mechanisms have also
been formerly suggested for the occurrence of core
bryophyte species on decaying logs (Söderström,
1989) and stream-dwelling angiosperms (Riis &
Sand-Jensen, 2002). Furthermore, local abundance
could also be positively related to larger-scale
 Distribution and abundance relationships in bryophytes 1881
distribution in stream bryophytes, although it would
then be unlikely that the mechanism driving the
pattern is related directly to metapopulation
dynamics. Rather, widely distributed and locally
abundant species might be generalists with regard to
environmental tolerance, while restricted and locally
rare specialist species would show the opposite trend.
Thus, if local abundance is also positively related to
distribution beyond a focal drainage system, then
processes other than local extinction and recolonisa-
tion are more likely to account for the pattern.
We tested these assumptions using occupancy–
abundance data from stream bryophyte species in a
boreal drainage basin. Rather than directly testing the
mechanistic basis of the positive distribution–abun-
dance relationship, we concentrated on examining if
abundance estimates of bryophytes measured from
local stream habitats correlated with: (i) their regional
occupancy (number of sites occupied in the focal
drainage system); (ii) provincial distribution in nor-
thern Europe; and (iii) global range size. We also
examined whether species characteristics account for
residual variation in the distribution–abundance rela-
tionship. Finally, similarly to the distribution–abun-
dance relationship, we examined distribution patterns
at two spatial scales: within the focal drainage system
and across northern Europe.
Methods
The focal study region comprises the Koutajoki
drainage basin in northeastern Finland (66–67N,
28–30E). The bedrock of the study area is highly
variable, with extensive areas of calcareous rocks.
Accompanied by considerable altitudinal differences,
this is mirrored in highly variable vegetation, ranging
from old-growth coniferous forests to nutrient-poor
bogs and fertile fens, as well as associated diverse
boreal flora. These factors also provide the basis for
the high variability of stream habitats across the
region. Headwater streams and small rivers in the
study area are characterised by circumneutral to
alkaline water, low turbidity and nutrient concentra-
tions ranging from low to moderate (Muotka &
Virtanen, 1995; Heino, Muotka & Paavola, 2003).
A set of 65 stream sites (first to fourth order, see e.g.
Allan, 1995) was surveyed for bryophytes in the
Koutajoki drainage basin. Bryophytes were collected
from eight or 10 50 cm · 50 cm quadrants allocated
 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd, Freshwater Biology, 51, 1879–1889
on a riffle site of c. 100 m2, following stratified random
or systematic sampling schemes. Both obligatorily
aquatic species (i.e. growing in permanently sub-
merged conditions) and semi-aquatic species (i.e.
those being only temporarily submerged or growing
on large boulders just above water level) were
recorded. Based on these surveys, regional occupancy
(i.e. the number of sites occupied in the Koutajoki
drainage basin) and mean local abundance (i.e. per-
centage cover over all quadrants at a site) at occupied
sites were calculated for each species. Further, the
distribution of the species found in the Koutajoki
survey was also estimated at two larger scales. These
were: (i) provincial distribution or the number of
provinces of the Nordic countries (Finland, Sweden,
Norway, Denmark and Iceland) occupied by each
species, according to Söderström (1996, 1998) for true
mosses and Söderström, Hassel & Weibull (2002) for
liverworts; and (ii) global distribution of species,
according to Dierßen (2001). Due to the rather coarse
level of information available for global distribution,
we generated five a priori distribution groups: (N)
species restricted to the Nordic countries (see above),
(E) European, (P) Palaearctic, (H) Holarctic and (G)
Global, i.e. occurring on both northern and southern
hemispheres. In practice, there were no N or E species
and only three P species, while those remaining were
either H or G. Therefore, we combined P and H species
in the same group, resulting in two distribution groups
for data analysis, Holarctic (H) and Global (G).
We tested the relationships between the distribution
and abundance of stream bryophytes using general
linear models. Our model included the dependent
variable of either regional occupancy (number of sites
occupied in the Koutajoki drainage system) or pro-
vincial distribution (number of provinces in northern
Europe occupied by the species in the Koutajoki
species pool), whereas explanatory variables included
mean local abundance (a continuous variable), major
species trait (aquatic/semi-aquatic, categorical) and
bryophyte group (true moss/liverwort, categorical).
We were interested primarily in the relationship
between the measures of distribution and mean local
abundance and, secondarily, in whether the two
categorical variables accounted for any of the residual
variation. The interaction term species trait · bryo-
phyte group was also included in the model. Further-
more, we examined whether the regional occupancy,
provincial distribution and mean local abundance of
1882 J. Heino and R. Virtanen
stream bryophytes were related to their global range cantly related to this measure of distribution (Table 2).
(H/G, categorical) using two-sample t-tests. Prior to Globally (G) distributed species were not more
analyses, regional occupancy (number of sites in the abundant locally (t ¼ 0.387, P ¼ 0.701) and did not
Koutajoki drainage basin) was log (x + 1) transformed occur at more sites in the Koutajoki basin than did
and mean local abundance (% cover) was arcsine- Holarctic (H) species (t ¼ )0.553, P ¼ 0.583). How-
squareroot transformed to improve normality and ever, G species occurred in significantly more prov-
remove heteroscedasticity. All analyses were run inces in northern Europe than H species (t ¼ 3.783,
using SPSS version 11.5 (SPSS Inc., 2002). P < 0.001).
There were marked differences in the species–sites
and species–provinces frequency distributions, the
Results
former which was related to regional occupancy and
We found a total of 47 aquatic and semi-aquatic the latter to provincial distribution. While most
bryophyte species from the 65 study sites in the species appeared to be rare with regard to their
Koutajoki drainage system (Appendix 1). Regional regional occupancy, that is, the frequency distribution
occupancy and mean local abundance of stream was strongly right-skewed (Fig. 2a), the same species
bryophytes were significantly related (all species showed almost contrasting frequency distribution
included: R2 ¼ 0.230, aquatic: R2 ¼ 0.359, P < 0.01, when their provincial distribution was considered
semi-aquatic: R2 ¼ 0.032, NS, Fig. 1), and significant
additional variation was related to species trait
(Table 1); that is, aquatic species occurred at signifi- Table 1 Results of general linear model for the relationships
cantly more sites than semi-aquatic species (Table 1, between regional occupancy (dependent variable) and mean
local abundance
Fig. 1). By contrast, provincial distribution was not
significantly related to mean local abundance, and Source d.f. MS F P
neither were the two categorical variables signifi-
Corrected model 4 0.594 4.872 0.003
Intercept 1 4.880 40.000 <0.001
Abundance 1 0.826 6.771 0.013
1.8 Group 1 0.087 0.709 0.404
tic Trait 1 0.580 4.755 0.035
ua
1.6 Aq Group · trait 1 0.256 2.095 0.155
Error 42 0.122
Regional occupancy (log sites + 1)

1.4 Categorical variables describing a major species trait (aquatic

versus semi-aquatic) and bryophyte group (true moss versus
1.2 liverwort) were included in the model to examine whether they
account for additional variation. Model R2 ¼ 0.317.
1.0
quatic
Semia Table 2 Results of general linear model for the relationships
0.8
between provincial distribution (dependent variable) and mean
local abundance
0.6
Source d.f. MS F P
0.4
Corrected model 4 246.019 0.929 0.456
Intercept 1 46 480.313 175.562 <0.001
0.2 Abundance 1 64.456 0.243 0.624
0 0.1 0.2 0.3 0.4 0.5 0.6 Group 1 79.658 0.301 0.586
Abundance (arsq cover) Trait 1 483.507 1.826 0.184
Group · trait 1 62.036 0.234 0.631
Fig. 1 The relationship between regional occupancy (log num- Error 42 264.752
ber of sites) and mean local abundance (arcsine-squareroot mean
cover). Aquatic species are denoted by filled squares and semi- Categorical variables describing a major species trait (aquatic
aquatic species by open squares. The solid regression line refers versus semi-aquatic) and bryophyte group (true moss versus
to the overall relationship between regional occupancy and local liverwort) were included in the model to examine whether they
abundance. account for additional variation. Model R2 ¼ 0.081.

 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd, Freshwater Biology, 51, 1879–1889

(a)
25
20
Number of species
15
10
5
0 0
0.02
Proportion of sites occupied
Fig. 2 Frequency distribution of species-proportion of sites (a) and species-proportion of provinces (b) occupied by stream bryophytes
of the Koutajoki flora.
(Fig. 2b). Thus, the stream bryophyte flora of the
Koutajoki drainage basin could be considered to be
characterised either by pronounced rarity or pro-
nounced commonness, depending on the measure of
distribution.
Discussion
The relationship between regional occupancy and
local abundance is important because it provides a
link between local population processes and regional
dynamics (Gaston & Blackburn, 2000; Freckleton et al.,
2005). The relationship can take several forms, but a
positive relationship is one of most common patterns
reported in the ecological literature (Gaston & Black-
burn, 2000). At best, such a known relationship can be
used to predict regional population size based on
mean local abundance or occupancy. The main
finding of this study was that the distribution–
abundance relationship depends on the measure of
distribution. Namely, local abundance correlated
significantly with the number of sites occupied in
the focal drainage basin, but it did not show a
significant relationship with the number of provinces
in which species were found. The former result agrees
with findings from recent studies of other stream-
dwelling organisms. For stream fishes, Tales et al.
(2004) found that local abundance showed a stronger
relationship with the number of sites occupied across
 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd, Freshwater Biology, 51, 1879–1889
Distribution and abundance relationships in bryophytes 1883
(b)
14
12
10
8
6
4
2
0.54 0.95 0.05 0.55 0.95
Proportion of provinces occupied
drainage basins than the number of river basins
occupied. Furthermore, for stream angiosperms, Riis
& Sand-Jensen (2002) found that the distribution–
abundance relationship was stronger within than
across drainage basins, and they suggested that
metapopulation dynamics explain the positive abun-
dance–occupancy relationship within a drainage sys-
tem (see also Freckleton & Watkinson, 2002).
According to metapopulation theory, high local
abundance decreases extinction probability and
increases emigration rate (e.g. Hanski, 1991). Riis &
Sand-Jensen (2002) reasoned that streams are char-
acterised by frequent disturbances which constantly
redistribute species via efficient passive dispersal
within a drainage system, and this process contributes
to a positive distribution–abundance relationship of
stream angiosperms. Although there is a dearth of
information about actual extinction rates, dispersal
rates and dispersal distances of stream bryophytes,
the physical characteristics of streams, frequent frag-
mentation of vegetative shoots, subsequent passive
dispersal of such fragments via flowing water and the
formation of bryophyte stands at suitable sites where
the fragments have settled (Stream Bryophyte Group,
1999; see also Johansson & Nilsson, 1993) all suggest a
potential role for similar mechanisms to that sug-
gested for stream angiosperms. Long-term observa-
tional data also show that many species of stream
bryophytes can successfully recolonise a fully
1884 J. Heino and R. Virtanen
denuded site in about 5–7 years, at least when the
source populations are located nearby (R.Virtanen &
T. Muotka, unpublished data).
We did not find a significant relationship between
local abundance and the larger-scale provincial dis-
tribution. If such a relationship existed, then the
mechanisms behind a positive distribution–abun-
dance relationship would be unlikely to be related to
metapopulation dynamics. Rather, for example,
widely distributed and locally abundant species
might typically utilise common resources or habitat
conditions available across a large region, while
restricted and locally rare species would show the
opposite (see also Hanski et al., 1993; Tales et al.,
2004). Further, alternative explanations for positive
distribution–abundance relationships at large scales,
as well as within a drainage system, include niche
breadth, range position and sampling artefact (see
Brown, 1984; Gaston et al., 1997). The lack of a
significant relationship between larger-scale distribu-
tion and local abundance may also indicate dispersal
limitation. In a recent modelling study, Freckleton
et al. (2005) suggested that in systems with low
colonisation rates, there are typically weak relation-
ships between distribution and abundance.
We found that the abundance–distribution relation-
ship differed between aquatic and semi-aquatic spe-
cies. Only aquatic species showed a significant
relationship between regional occupancy and local
abundance, whereas no significant relationship was
found for semi-aquatic species. This result contrasts
with a previously reported finding that obligatorily
submerged plants had a lower coefficient of deter-
mination of the abundance–occupancy relationship
than amphibious plants (Riis & Sand-Jensen, 2002).
Riis & Sand-Jensen (2002) suggested that their finding
resulted from amphibious plants being more effec-
tively dispersed than obligatory aquatic plants,
because the former may utilise well both aerial seed
dispersal and hydrochory. For stream bryophytes that
presumably rarely rely on spores and shoot fragments
for dispersal and site colonisation, differences
between aquatic and semi-aquatic species in aerial
dispersal traits may not be that profound. Rather, we
suggest that aquatic bryophyte species are, on
average, more widely distributed and show a stronger
distribution–abundance relationship within a stream
system than semi-aquatic species, because the shoot
fragments of the former group are more closely
 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd, Freshwater Biology, 51, 1879–1889
associated with and more efficiently distributed by
the main medium of dispersal, that is, flowing water,
as well as organisms potentially contributing to
dispersal (Glime & Vitt, 1984). Obviously, this reason-
ing cannot be extended fully to patterns across stream
systems, as dispersal between even neighbouring
catchments is obviously only likely to happen via
airborne spores or via zoochory. Alternatively, many
aquatic bryophytes have flexible morphology, and
they can thus occur in a wide variety of microhabitats
(Suren, 1996) and obviously in different types of
streams, which might contribute to their overall
distribution patterns. Thus, aquatic species may also
have a larger niche breadth than semi-aquatic species.
However, the degree to which aquatic and semi-
aquatic bryophytes differ in their morphological
flexibility and niche breadth is poorly known, and
such differences are very speculative.
An issue related to the abundance–distribution
relationship is the pattern of frequency distribution
of species (e.g. Hanski, 1982; Collins & Glenn, 1997).
We found that the pattern differed with regard to the
scale of distribution (i.e. regional occupancy versus
provincial distribution) and type of distributional
data. Although such a clear difference between the
species-regional occupancy and species-provinces
frequency distributions might seem unexpected, this
finding is by no means new, but such scale-depend-
ence was noted in studies on the distribution
patterns of plants decades ago (see McIntosh, 1962
and references therein). Thus, there are plausible
reasons to expect that it might be a rather general
pattern. For instance, it is likely that there are at least
a few suitable sites for each species in large-scale
provinces or geographical grids, and thus most
species would occur in several or even most of such
large-scale units. By contrast, within a single region
or a drainage system, most species are similarly able
to occur at only a few sites and a minority of species
at many sites, and this would lead to a right-skewed
frequency distribution of regional occupancy, with
most species being rare. It follows that the Koutajoki
stream bryophyte flora can be characterised by either
pronounced rarity or pronounced commonness,
depending on the measure of distribution (Fig. 2).
Thus, the patterns detected may not only be affected
by spatial extent (e.g. Collins & Glenn, 1997; Bossuyt
et al., 2004), but also strongly affected by grain size
(e.g. Guo et al., 2000), as well as the combination of
 Distribution and abundance relationships in bryophytes 1885
these two aspects of spatial scale (Wiens, 1989; see species, correspondingly (Gibson et al., 1999). In our
also McGeoch & Gaston, 2002). However, it is worth data on the regional occupancy of stream bryophytes,
noting that the species-provinces frequency distribu- true core species were certainly lacking, and the most
tion incorporating all stream bryophyte species in widely distributed species occurred at a little more
northern Europe, for example, would look rather than 60% of sites. However, a number of species
different from that we found for the Koutajoki stream groups with regard to their regional distribution and
bryophyte flora. Thus, it appears that the association local abundance could be identified (Fig. 3). For
between species pool, spatial extent and grain size is example, (i) there were large species (e.g. Fontinalis
also important when distribution patterns are con- antipyretica, Fontinalis dalecarlica, Hygrohypnum ochrac-
sidered. eum) that were relatively abundant locally and widely
Patterns in species’ regional frequency distributions distributed. Thus, these fontinaloid-type species rep-
have typically been considered in the context of core resent the closest candidates for core or dominant
and satellite species hypothesis (CSS, Hanski, 1982) bryophyte species in boreal streams, although they
and, more recently, in terms of Grime’s (1998) classi- appear to be absent from a large proportion of sites.
fication of dominant, subordinate and transient spe- Additionally, (ii) there were two species (Brachythe-
cies (DST, Gibson et al., 1999). Both of these concepts cium rivulare, Leptodictyum riparium) that had rather
divide species into three groups based on their restricted regional distribution, but were locally
occurrence across sites within a region: in the CSS abundant, at least at some sites. Furthermore, (iii)
there are core species (i.e. those occurring at most there were species that had low cover locally, but
sites), satellite species (i.e. those occurring at few sites) were relatively widely distributed. These might be
and intermediate species, and in the DST of interest considered as intermediate species, and combine
are dominant, transient species and subordinate different tactics of subordinate species (sensu Grime,

1.8

FISPUS FONANT
1.6
BLIACU
BRYPSE
CHIRIV
1.4 SCAUND
FONDAL
PALFALC
Regional occupancy (log sites + 1)

JUNPUM

HYGORC
1.2 SCHAGA HYGALP
CALGIG JUNCOR PLARIP
HYGSMI
FISOSM CAMELO
1.0 CHIPOL HYGFLU

PHIFON DICPEL HYGLUR

FISADI
BRARIV LEPRIP
0.8
BRAPLU HYGDUR
PELLIA
Fig. 3 Differences between stream bryo- LEIBAN
phyte species in the relationship between 0.6
regional occupancy (log number of SCOSCO CRAFIL DICFAL
sites + 1) and mean local abundance MARCH
(arcsine-squareroot mean cover). Aquatic WARTRI SCOREV LEIGIL
0.4
species are denoted by filled squares and TAYLIN DICPAL PALCOM
semi-aquatic species by open squares. WAREXA
PHITOM PHICAL
Species are denoted by the first three let-
ters of genus and species names (Appen- 0.2
dix 1). For clarity, acronyms of some 0 0.1 0.2 0.3 0.4 0.5 0.6
species with low local abundance and
restricted distribution are not shown. Abundance (arsq cover)

 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd, Freshwater Biology, 51, 1879–1889
1886 J. Heino and R. Virtanen
1998). In the life-history scheme of Muotka & Virtanen
(1995) these species belong to the Leptodictym-Bryum
type. Lastly, (iv) there were species with low
abundance and restricted distribution, clearly belong-
ing to the group of satellite species or transient species
(Fig. 3).
Our findings nevertheless deviate from those of
previous workers who have found frequency distri-
butions that are typified either by well-defined core
and satellite groups or, alternatively, have a long
positive tail, with at least some species that occur at
most sites (Hanski, 1982; Collins & Glenn, 1997; Guo
et al., 2000). Furthermore, it has been found that
bryophytes on forest logs fit relatively well with the
CSS hypothesis, with distinctive groups of core and
satellite species (Söderström, 1989). The degree to
which, and the reason why, stream bryophytes
differed from the patterns typical of many other
organism groups is presently uncertain, but a possible
reason is again related to the characteristics of
streams. For example, one might envisage that the
pronounced spatial heterogeneity of stream habitats
(e.g. large boulders are absent from some sites and
thus there is no habitat for most semi-aquatic bryo-
phytes) clearly limits the occurrence of species.
Alternatively, as streams are characterised by fre-
quent and often drastic disturbances, such as flash
floods and ice scour in boreal regions, even the most
dispersive bryophyte species may be temporarily
absent from a high proportion of potential growing
sites. If this reasoning is correct, however, then one
could also expect rather similar distribution patterns
for other stream-dwelling organism groups. This is
not exactly the case as, for both stream insects (Heino,
2005) and stream diatoms (Soininen & Heino, 2005),
the frequency distributions of regional occupancy do
not have a distinct core species group, are charac-
terised by numerous satellite species, yet they still
have a long positive tail with at least one ubiquitous
species. However, one could also invoke the ecologi-
cal differences between stream bryophytes, insects
and diatoms to account for their differing distribution
patterns. For instance, growing firmly attached to
stone surfaces, stream bryophytes are: (i) probably the
most disturbance prone of these three organism
groups; and (ii) the slowest to colonise sites after
disturbance. Yet, even these environmental conditions
and organismal characteristics of stream bryophytes
cannot fully account for the atypical form of the
 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd, Freshwater Biology, 51, 1879–1889
species-occupancy frequency distribution, as a wide
variety of patterns have also been found in other
systems (e.g. Collins & Glenn, 1997; Bossuyt et al.,
2004). However, we suggest that environmental con-
ditions in terms of disturbance frequency and severity
may potentially contribute to the frequency distribu-
tion of species, as well as the strength of the
distribution–abundance relationship.
Acknowledgments
This study was supported by grants from the Acad-
emy of Finland (to JH) and the Ministry of Environ-
ment in Finland (to RV). We thank Alan Hildrew and
anonymous referees for valuable comments on earlier
versions of the manuscript.
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 Distribution and abundance relationships in bryophytes 1889
Appendix 1 List of aquatic and semi-aquatic bryophyte species found in the Koutajoki drainage basin in northeastern Finland

Species Group Trait Cover Range Provinces Occupancy

Amblystegium fluviatile (Hedw.) Schimp. T A 4.01 G 52 8

Blindia acuta (Hedw.) Bruch & Schimp. T S 2.43 G 75 29
Brachythecium plumosum (Hedw.) Schimp. T S 0.52 G 79 4
Brachythecium rivulare Schimp. T S 15.96 G 77 5
Bryum pseudotriquetrum (Hedw.) P. Gaertn. et al. T S 0.81 G 79 26
Calliergon cordifolium (Hedw.) Kindb. T S 0.70 G 87 2
Calliergon giganteum (Schimp.) Kindb. T S 1.87 H 82 11
Calliergonella lindbergii (Mitt.) Hedenäs T S 0.69 G 75 2
Campyliadelphus elodes (Lindb.) Kanda T S 1.54 P 40 10
Chiloscyphus polyanthus (L.) Corda L S 0.52 H 78 9
Chiloscyphus pol. var. rivularis (Schrad.) Gottsche et al. L A 7.67 H 32 24
Cratoneuron filicinum (Hedw.) Spruce T S 1.54 G 77 2
Dichelyma falcatum (Hedw.) Myrin T S 2.52 H 65 2
Dichodontium pellucidum (Hedw.) Schimp. T S 0.37 H 72 6
Dicranella palustris (Dicks.) Crundw. ex E.F. Warb. T S 0.40 H 62 1
Fissidens adianthoides Hedw. T S 0.69 G 87 6
Fissidens osmundoides Hedw. T S 0.42 G 83 9
Fissidens pusillus (Wilson) Milde T A 1.34 H 33 39
Fontinalis antipyretica Hedw. T A 13.08 G 83 40
Fontinalis dalecarlica Bruch & Schimp. T A 18.60 H 67 21
Hygrohypnum alpestre (Sw. ex Hedw.) Loeske T S 4.67 H 56 13
Hygrohypnum duriusculum (De Not.) D.W. Jamieson T A 1.77 H 57 4
Hygrohypnum luridum (Hedw.) Jenn. T S 3.75 G 71 7
Hygrohypnum ochraceum (Turner ex Wilson) Loeske T A 18.52 H 76 14
Hygrohypnum smithii (Sw. ex Lilj.) Broth. T A 9.05 H 33 10
Jungermannia exsertifolia ssp. cordifolia (Dumort.) Vána L A 6.22 H 49 11
Jungermannia pumila With. L A 8.27 G 62 19
Leiocolea bantriensis (Hook.) Jörg. L S 0.35 H 53 3
Leiocolea gillmanii (Austin) A. Evans L S 2.70 H 50 2
Leptodictyum riparium (Hedw.) Warnst. T S 25.01 G 59 5
Marchantia polymorpha L. L S 4.85 G 84 2
Oncophorus wahlenbergii Brid. T S 0.25 H 78 1
Palustriella commutata (Hedw.) Ochyra T S 2.50 G 53 1
Palustriella falcata (Brid.) Hedenäs T A 9.80 P 65 20
Pellia epiphylla (L.) Corda + P. neesiana (Gottsche) Limpr. L S 2.62 H 83 4
Philonotis calcarea (Bruch & Schimp.) Schimp. T S 1.25 H 49 1
Philonotis fontana (Hedw.) Brid. T S 0.26 G 84 6
Philonotis tomentella Molendo T S 0.03 H 67 1
Plagiomnium ellipticum (Brid.) T.J. Kop. T S 0.10 G 82 1
Rhynchostegium riparioides (Hedw.) Cardot T A 19.21 G 54 12
Scapania undulata (L.) Dumort. L A 4.00 H 83 23
Schistidium agassizii Sull. & Lesq. T S 1.24 H 68 13
Scorpidium revolvens (Sw. ex Anonymus) Rubers T A 0.97 G 80 2
Scorpidium scorpioides (Hedw.) Limpr. T A 0.03 G 88 2
Tayloria lingulata (Dicks.) Lindb. T S 0.25 H 45 1
Warnstorfia exannulata (W. Gümbel) Loeske T A 3.13 G 86 1
Warnstorfia trichophylla (Warnst.) Tuom. & T.J. Kop. T A 0.38 H 45 2

Group: true mosses (T) and liverworts (L). Trait: obligatory aquatic (A) and semi-aquatic (S). Cover: mean percentage cover at
occupied sites. Range: global (G), Holarctic (H) and Palaearctic (P). Also shown are the number of provinces (n ¼ 91) and sites (n ¼ 65)
occupied by each species of the Koutajoki bryophyte flora.

 2006 The Authors, Journal compilation  2006 Blackwell Publishing Ltd, Freshwater Biology, 51, 1879–1889