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Wind dispersal of spores with focus on bryophytes
Niklas Lönnell
Plants & Ecology Plant Ecology 2011/3 Department of Botany Stockholm University
Wind dispersal of spores
with focus on bryophytes
by
Niklas Lönnell
Supervisors: Kristoffer Hylander, Johan Ehrlén, Sebastian Sundberg,
Bengt Gunnar Jonsson
Plants & Ecology
Plant Ecology 2011/3 Department of Botany Stockholm University
Plants & Ecology
Plant Ecology
Department of Botany
Stockholm University
S-106 91 Stockholm
Sweden
© Plant Ecology
ISSN 1651-9248
Printed by FMV Printcenter
Cover: The fungi Phaeolus schweinitzii, the moss Funaria hygrometrica, and the fern
Botrychium boreale. Photographs: Niklas Lönnell
3
Summary
Investigation of dispersal, especially long-distance dispersal (LDD), is of utmost importance for
the understanding of many patterns and processes like species distributions, range expansions
including responses to climate change and metapopulation dynamics. The exact distance beyond
which dispersal can be defined as LDD is dependent on the species and can vary from 100
metres to intercontinental distances. The wind dispersal of small diaspores has been poorly
studied (Diaspore sizes: e.g. in bryophytes (diameter 10–50(–100) μm), fungi (3–10–(300) μm),
ferns (30–100 μm). According to Stoke's law they can travel farther than larger diaspores of the
same shape and density given the same wind speed, even if larger seeds can increase the
buoyancy with features like pappi or wings. Diaspore size, shape, weight, type of abscission
(active or passive), abscission height, abscission time, horizontal wind speed and thermal
updrafts/turbulence can influence the dispersal distances. The source i.e. the number of diaspores
produced is very strong in many species (cfr the production of 100 million spores/m2
in
Pleurozium schreberi). The most common direct method for studying dispersal includes putting
out a point source and a number of spore traps at increasing distances. Indirect methods
encompass approaches such as comparing 1. the genetic similarity between populations in
different parts of a country or continent, 2. successful colonizations with connectivity (i.e.
studying the patterns of realized dispersal), 3. the longevity of diaspores with the species
distribution and 4. the species composition with prevailing winds. The question concerning the
temporal and spatial scales on which species with small diaspores are dispersal limited still
remains to be answered.
Sammanfattning
Kunskapen om hur arter sprider sig särskilt över långa avstånd är viktiga för att förstå t.ex. arters
utbredningsmönster, förändringar i artutbredningar som svar på t.ex. klimatförändringar. Det
exakta avståndet över vilket spridning räknas som räknas som långdistansspridning kan variera
från 100 m till avstånd mellan olika kontinenter. Vindspridning av små spridningsenheter har
inte studerats i någon större utsträckning. (Storleksexempel: mossor (diameter 10–50(–100) μm),
svampar (3–10–(300) μm), ormbunkar (30–100 μm)). De kan sprida sig längre än större frön
som dock ibland har olika slags anpassningar som hårpenslar och vingar för att kunna spridas
längre. Spridningsenhetens storlek, form, vikt samt hur (aktivt eller passivt) och på vilken höjd
de kommer ut i luften, vindhastighet och om det förekommer turbulens eller varmluft som stiger
kan påverka spridningsavstånden. Antalet spridningsenheter som produceras är stort hos många
arter (t.ex. 100 miljoner sporer per kvadratmeter hos väggmossa Pleurozium schreberi). En
vanlig metod att studera spridning är sätta ut en källa och ett antal fällor på olika avstånd. Även
indirekta metoder använts så som att jämföra 1. genetisk skillnad mellan olika lokaler och
populationer 2. framgångsrik etablering med hur långt det är till närmaste möjliga källor 3. hur
långlivade spridningsenheterna är och vilken utbredning arten har samt 4. artsammansättning
med förhärskande vindriktningar. I vilken utsträckning i fråga om tid och rum som arter med små
spridningsenheter är begränsade av sin spridningsförmåga återstår att utreda.
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Contents
Summary ........................................................................................................................................................................................... 3
Sammanfattning ................................................................................................................................................................................ 3
Contents ............................................................................................................................................................................................ 4
What is dispersal? ............................................................................................................................................................................. 5
Transportation ................................................................................................................................................................................... 6
Abscission ......................................................................................................................................................................................... 7
Abscission height .............................................................................................................................................................................. 9
Buoyancy ........................................................................................................................................................................................ 10
Diaspore shape ................................................................................................................................................................................ 11
Diaspore size ................................................................................................................................................................................... 11
Diaspore longevity .......................................................................................................................................................................... 15
Number of diaspores ....................................................................................................................................................................... 16
Diaspore banks ............................................................................................................................................................................... 17
Dormancy ....................................................................................................................................................................................... 19
Deposition ....................................................................................................................................................................................... 19
How to study dispersal .................................................................................................................................................................... 20
Direct methods ........................................................................................................................................................................... 20
Lagrangian methods – tracking individuals/single diaspores ............................................................................................... 20
Eulerian methods – trapping ................................................................................................................................................ 21
Indirect methods ........................................................................................................................................................................ 23
Biogeographical studies ........................................................................................................................................................ 26
Other studies ......................................................................................................................................................................... 27
Models ....................................................................................................................................................................................... 28
Phenomenological models..................................................................................................................................................... 28
Mechanistic models ............................................................................................................................................................... 29
Conclusions .................................................................................................................................................................................... 29
References ...................................................................................................................................................................................... 30
5
What is dispersal?
The general meaning of the word "dispersal" can be stated as "the action or process of
distributing or spreading things or people over a wide area" (Soanes & Stevenson 2005). In a
biological context definitions like "intergenerational movement" (Bullock et al. 2002), "the
actual scattering or distributing of organisms on earth's surface"; "transport of diaspores"
(Holmes 1979) or "unidirectional movement of an individual away from its place of birth"
(Clobert et al. 2001; Nathan 2008) may be more appropriate. As Pijl (1982) remarks, the term
dispersal has been used synonymously with distribution which may lead to confusion when
reading older publications. Since this work will be restricted to plants, a definition relevant for
this context could be "transportation of diaspores away from the plant of origin". "Diaspore" is
general term for a plant dispersal unit for example a seed, a spore, or a gemma.
The term "long-distance dispersal" (LDD) is more difficult to define. How far is long? The
absolute distance varies between different organism groups as well as species. There are two
ways of defining the approximate distance beyond which dispersal can be defined as LDD: to set
an absolute value or to set a relative value beyond which a certain low percentage of diaspores
are being dispersed. Cain et al. (2000) suggest 100 m as an absolute value and a relative value
based on the distance beyond 1 % of the diaspores travel. In the examples given by Nathan
(2008) the distance is a few hundred metres. Nathan (2005) suggests one should write the most
relevant value for the study in question first, followed by the corresponding value for the other in
parentheses after e.g. 1000 m (0.1 %). For Sphagnum Sundberg (2005) defines LDD as dispersal
more than 1 km which is beyond the normal-sized mire in the study area. However, when
Zanten & Pócs (1981) discuss "long-range dispersal" only distances over 2000 km were
considered.
l
Production
Dis
per
sal
Abscission
Transportation
Deposition
Establishment
Figure 1. The different stages of dispersal. Realized or effective dispersal can include establishment as
well as production of new diaspores.
6
The dispersal can be divided into three different stages: abscission (release, liberation and
discharge), transport (transportation, transfer) and deposition (capture) (Fig.1). But for dispersal
to have an effect, post-dispersal events like germination and survival of the progeny must be
successful. Hence "Effective dispersal" includes the establishment of a new plant that can
reproduce itself (Nathan 2008). This can also be named "realized dispersal". At the regional
scale species can be principally dispersal limited or habitat limited (Münzbergová & Herben
2005). The establishment phase is critical for many bryophytes and both germination and
protonema growth could be limited by factors such as pH and water availability (Wiklund &
Rydin 2004). As the production (formation) of diaspores is also a vital component in the success
of dispersal, it may be included in an extended definition of what successful dispersal
incorporates (Löbel et al. 2009).
I will deal with dispersal in space in this paper. Dispersal in time is, however, briefly discussed
under the title "Diaspore banks".
Transportation
The term used to describe the transportation is often a combination of a prefix describing the
name of the agent in Greek and a suffix describing what kind of object that is being transported.
Pijl (1982) describes the terminology extensively and I summarise it as follows. The suffix "-
gamy" can be used for describing the transportation of and differentiation of gametes. The suffix
"-phily" can be used of the transport of spores, especially microspores and their successors,
pollen grains. The suffix implies that the transportation is to a friendly place e.g. the stigma/the
megaspore (philein (Gr.) = to love). The suffix "-chory" is used for mere dispersal of seeds etc.
(chorein (Gr.) = to wander). When referring to the transportation of isospores (same spore size in
both sexes) of cryptogams both "-phily" and "-chory" can be used.
I will use "-chory" for transportation of spores in bryophytes and fungi.
Dispersal by an animal vector may expand the range as well as improve the percentage of
diaspores that end up on a suitable substrate. This paper, however, will deal with abiotic vectors,
7
more precisely wind dispersal of spores (anemochory) with some references to wind dispersal of
seeds (anemochory) and wind pollination (anemophily).
Bryophytes are generally considered to be hydrogamous even if Cronberg et al. (2006)
demonstrated that springtails and mites may act as vectors of bryophyte gametes as well.
Abscission
There are a multitude of ways and adaptions of how to release the diaspores into the air.
Abscission can be active and explosive or passively triggered by wind, convection or raindrops.
The role of abscission timing has been found to be an important factor for LDD (Greene 2005;
Skarpaas et al. 2006; Soons & Bullock 2008).
Vascular plants
Pollen in wind-dispersed species is mostly released passively. In many species with dust seeds
the abscission is passive and the seeds are discharged as the wind moves the drying capsule.
In Polypodiaceae the spores are actively discharged when the thickened sporangium wall is
thrown back as the water content in the cells decreases. However in other species within
Marattiales and Ophioglossales the discharge is passive. Club mosses Lycopodium-species have
passive discharge while Selaginella-species have active. (Ingold 1974)
Liverworts
Most leafy liverworts have a capsule which opens entirely when the capsule wall breaks along
four lines which expose the spore mass when it dries out. Inside the capsules there are elaters, up
to 0.5 mm long tubes with spiral reinforcement in the walls. In some species (e.g. Cephalozia
bicuspidata) the elaters spring loose from the capsule wall when they dry and hence can bring
spores up in the air. In others, the elaters show more moderate hygroscopic movement (e.g.
Marchantia polymorpha) and in still others they show very slight hygroscopic movement (e.g.
Pellia epiphylla). In some species (e.g. Frullania dilatata) both ends of the elaters are fixed in
the upper and lower part of the capsule wall and they act as spiral springs and cause a violent
opening of the capsule which can throw spores 2 cm up in the air. (Ingold 1974)
8
Mosses
Unlike in liverworts, abscission is passive in most moss species. However in Sphagnum the
spores are released explosively up to 20 centimetres in the air when the lid of the capsule
dehisces (Ingold 1974). Turbulent vortex rings are created and facilitate dispersal of the spores
(Whitaker & Edwards 2010). How high the spores reach varies according to capsule size
(Sundberg 2010b). The violent abscission of the spores is due to air pressure at the basal part the
capsule under the spore sack and has been compared to firing an air gun (Ingold 1974). That the
air pressure causes the eruption has been questioned (Duckett et al. 2009) and the discussion
continues about what causes the violent abscission (Duckett et al. 2010; Sundberg 2010a). The
capsule of Andreaceae opens in four narrow openings along the side of the capsule. The top of
the capsule is still intact unlike the case in liverworts. In the Polytrichaceae the capsule has a
peristome which is like a membrane with small holes through which spores can escape, when
wind moves the capsule, like from a saltcellar.
Within the true mosses (Bryopsida) most species have a capsule that dehisces with a lid and the
mouth of the capsule has a peristome with one or two layers of teeth. The peristome teeth move
when air humidity changes (hygroscopic). In the vast majority of moss species the peristome
shuts when moist and opens when it dries out (xerocastique). However in a few species the
peristome opens when it is moist (hygrocastique) (Hedenäs 1992). Hygrocastique peristomes
occur foremost in epiphytic species in e.g. the families Meteoriaceae and Orthotrichaceae
(Zygodon, Macromitrium, Orthotrichum, Orthomitrium) and the genera Pylaisia, Leucodon,
Cryphaea and in the species Coscinodon cribosus (cfr. Hedenäs 1992; Porley & Hodgettts
2005).
The hygroscopic movement of the outer peristome teeth can throw spores into the air. During
171 such movements the number of discharged spores was calculated to be 15,000 out of a total
of approximately 500,000 spores in one capsule of Eurhynchium confertum (Ingold 1974). The
peristome can regulate when the spores are to be released and the peristome structure can also
affect wind conditions and cause turbulence which can facilitate spores to become airborne.
During dry conditions the probability of convection and longer transportation are higher. In wet
conditions deposition near the capsule of origin is higher through wet deposition. So why do
9
some species have a hygrocastique peristome that prevents dispersal during dry conditions?
Many of those species are epiphytes and the establishment phase is suspected to be the
bottleneck for many epiphytic species (Löbel 2009). A higher establishment rate may be more
important than long dispersal in these species. But some epiphytes as well as species on other
substrates lack a peristome or have a heavily reduced one. Why is that? Vitt (1981) implies that
this is a character of xerophytic species, but he offers no explanation for the benefit of this in a
dry environment. Shaw & Robinson (1984) note that many species with erect capsules lack a
peristome, but they also remark that as many of the species with erect capsules, e.g. epiphytes,
grow on perpendicular surfaces and the orientation of the capsules becomes in fact horizontal
and within the genus Schizymenium the species with the most advanced peristome have erect
capsules whereas the ones with reduced peristome are pendant.
Raindrops can also trigger spores to leave the capsule, and the splash cups with gemmae in the
thallous liverwort Marchantia polymorpha extend gemma transportation distances.
Fungi and Lichens
In fungi and lichens the spores are released both by active and passive discharge. Some
ascomycetes generate a wind by synchronizing the abscission of spores (Roper et al. 2010). In
many polypores the discharge away from the fruiting body is passive where the spores fall from
console-shaped fruitbodies even if the release from the asci is active.
Abscission height
The height from which the diaspores are released influences how far the diaspores travel. Firstly,
it influences the time for the diaspore to reach the ground, which increases the chance that
updrafts or winds will move the diaspore vertically and horizontally. Secondly, the amount of
time that the abscission point is above the laminar layer is influenced and hence the opportunities
for diaspores to be released in wind conditions more suitable for dispersal.
If thermal updrafts are disregarded the distance (xd) that a diaspore travel can be described as a
function of the abscission height (h) and horizontal wind velocity (u) and the terminal velocity of
the diaspore (vt) (Kuparinen 2006):
10
xd = hu/vt
In mosses it ranges from less than one millimetre to tens of centimetres for those species with the
longest setae. In liverworts it ranges from Riccia whose spores are released only when the thallus
is disintegrated to e.g. Marchantia with up to 1 decimetre tall gametophores. In the average leafy
liverwort the thin, hyaline seta is elongated to a few centimetres after the spore capsule is fully
developed. In club mosses as well as horsetails Equisetum spp. the spores are released at the
highest point of the plant which range from a few centimetres to over a metre in the tallest
species. Ferns release spores from specialized fronds or from the sporangia on the lower side of
unspecialized fronds and the height varies along with the height of the species from a few
centimetres to several metres in tree ferns. Many basidiomycetes release spores from the
underside of the hat of the fruit body toadstool/mushrooms. The abscission height in ground
dwelling lichen is rather small as well, from millimetres to a few centimetres.
Many epiphytic or epilithic species reaches much higher and the abscission heights for these
species are noticeably increased. If there is a dense canopy the wind speed may be negatively
affected.
Buoyancy
Buoyancy is sometimes used to describe the ability of a diaspore to float in water. However, in
the context of this report it is used to describe the ability for a diaspore to "float" or remain in the
air. This ability is often estimated by the terminal velocity (=falling velocity or settling speed).
The terminal velocity can be estimated from Stoke's law. It applies to spherical objects with a
smooth surface. For very small and very large diaspores one can apply some corrections to adjust
the equation to better reflect the true terminal velocity (Gregory 1973). This approximation must
be complemented with measurements to determine the true terminal velocity that can deviate
from the ideal terminal velocity due to deviating (non-spherical) shape, surface structures and
air-filled spaces within the diaspore. The terminal velocities for pollen are from 2 to 6 cm/s
(Whitehead 1983).
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Stoke's law: vs = (2/9) (σ-ρ/μ) *gr2
vs = terminal velocity
σ = density of sphere in g/cm3 (water = 1.00)
ρ= density of medium (air = 1.27x10-3
g/ cm3)
g = acceleration of gravity (981 cm/s2)
μ = viscosity of medium (air at 18° C = 1.8 x 10-4
g/ cm/s)
r =radius of sphere in cm
There are also simplifications to similar estimates for non-spherical objects (Sundberg 2010b).
Diaspore shape
The shape varies much among different spores and pollen types. Many are rounded but others are
cylindrical, oval, kubic etc. Most bryophyte spores are more or less isodiametric while fungal
spores vary more in shape by often being longer than wide.
Among the 303 european polypores investigated by Kauserud et al. (2008) most species have a
spore shape ratio between 1 and 3. The largest ratio (6.5) belonged to Skeletokutis stellae. White
rot species and parasites have more spherical spores than brown rot species and saprotrophs, and
larger spores tend to be more spherical.
How the shape affects the terminal velocity and hence the dispersal capacity is not fully
illuminated. It has been suggested, however, that an elongated shape should decrease the
terminal velocity (McCubbin 1944; Gregory 1973).
Diaspore size
In different taxonomical groups and different publication the size of a diaspore can be estimated
in length and width, diameter, area (length x width), volume and weight. To be able to compare
the different estimates I have made the following conversion table (Table 1). However this is a
great simplification since it assumes that the diaspore is smooth, spherical and has the density of
1 g/cm3.
12
As the size increases the terminal velocity increases. What exact implication this has for
dispersal is harder to predict. In aerosols (when particles are suspended in air like smoke) most
particles have a diameter less than 5 μm. Wilkinson (2001) investigated testate amoeba
assemblages in the Artic and Anarctic and found that the largest species confined to one of those
areas was 230 μm while those with a cosmopolitan distribution had a maximum size of 135 μm.
Table 1. Overview over different measures used to define the size of diaspores. The conversion
anticipates that the diaspore is smooth, sphaerical and has a specific density of 1 g/cm3.
Diameter Area Volume Weight Terminal velocity
μm μm2 μm
3 mg mm/s
1 1 0.5 0.0005 0.03
5 25 65 0.07 0.76
10 100 523 0.52 3
20 400 4187 4 12
50 2500 65417 65 76
100 10000 523333 523 302
200 40000 4186667 4187 1210
500 250000 65416667 65417 7560
Pollen grains and spores vary much in size; From 1–5 μm (spores of Penicillum and Aspergillus
and pollen of Myosotis) to 200 μm (e.g. pollen in Curbitaceae and Nyctaginaceae) or even 800
μm (megaspores in Selaginella). (Nilsson 1980). However, most wind-dispersed spores and
pollen are usually smaller.
Pollen grains of most anemophilous plants have a diameter of 20–30(–60) μm. Some species
have pollen grains with air sacs to facilitate the dispersal. One example of this is pines Pinus spp.
that have large (50–150 μm) and heavy pollen grains (Faegri & Pijl 1979).
Pijl (1982) mentions examples of weights from some families with dust diaspores: 0.003 mg
(Orchidaceae), 0.004 mg (Pyrolaceae) and 0.001 mg (Orobanchaceae). The seeds from the
13
Histogram of spore size in liverworts
Mean spore diameter (µm)
Nu
mb
er
of sp
ecie
s
20 40 60 80 100 120
01
02
03
04
05
0
Finnish species of Pyrolaceae are 200 μm to 1 mm long and 80 μm to 200 μm broad (Pyykkö
1968). The seeds of Orchidaceae are also elongated and some examples of seed lengths in
Orchidaceae range from 0.19 to 3.8 mm (Clifford & Smith 1969). The larger orchid seeds may
thus not qualify as dust seeds. Seeds that are lighter than 0.02 mg–0.05 mg and seeds with
plumes and wings can be classified as having wind as the primary dispersal vector (Greene &
Calogeropoulos 2002).
Ferns tend to have larger spores than mosses and have higher points of initial spore release
(Ingold 1974). Spores of ferns are 30–100 μm (Tryon & Lugardon 1990 in Wolf et al. 2001).
Spores of Lycopodium and Selaginella in Taiwan range from 20–40 μm (Huang 1981).
Histogram of spore size in mosses
Mean spore diameter (µm)
Nu
mb
er
of sp
ecie
s
50 100 150 200
02
04
06
08
01
00
12
0
Figure 2. The mean spore size of 1035 British bryophyte species according to Hill et al. (2007).
Most bryophyte spores have a diameter that is less than 50 μm. A higher proportion of the
liverworts than of the mosses have spore diameters in the range from 40 to 100 μm (Hill et al.
2007). (Fig. 2)
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The largest moss spores are found in Archidium. Most of the species in this genus have an
average spore diameter between 100 μm and 200 μm. The species with the largest average spore
diameter is A. brevinerve 235 (200–260) μm whereas the one with the largest measured spore is
A. ohioense 162 (110–310) μm (Snider 1975). The species that is widespread in Europe is A.
alternifolium 180 (139–223) μm (Boros et al. 1993). The cleistocarpous capsule has a very short
seta and is hidden among the leaves. The spores are spread when the capsule wall is ruptured by
decomposition. This suggests that wind dispersal is not likely in Archidium and other vectors
should may be important, e.g. water. The same could be said about the species in the thallous
liverwort genus Riccia which also have large spores.
As Mogensen (1981) suggests the spore diameter distribution in a single species may not be
unimodal. The bimodality can be due to that some spores are aborted or that the sexes have
spores of different size. It may be a matter of resolution and accuracy of measurement as he
showed for Ceratodon purpureus.
The main focus in this report has been on spores as they usually are smaller and hence more
likely to be contributing to the LDD of most species. However, it is important not to forget that
bryophytes and lichens can disperse by means of fragmentation as well as by specialized asexual
diaspores. Fragments and asexual diaspores have an advantage in the establishment phase and
are generally thought to contribute to short-distance dispersal. The fragments of bryophytes
retrieved from a late snowbed in early spring in the alpine zone consisted primarily of species
occurring in the vicinity, but a few species from the subalpine belt were found as well (McDaniel
& Miller 2000). This implies that in open habitats fragments can even be dispersed by wind over
the snow but it does not contradict that this includes mostly short-distance dispersal. However, in
Anastrophyllum hellerianum the spores and gemmae are of equal small and are thought to be
able to contribute to LDD as well (Pohjamo et al. 2006). Tubers seem unlikely to disperse by
wind since they mostly develop on rhizoids in the soil. Lichens have a diversity of specialized
modes for asexual propagation. Lichhen soredia were the most common diaspores in a survey
with Rotorod samplers on Signy Island in Antarctica (Marshall 1996).
Most fungal spores have a diameter of around 10 μm but they range from 3 μm in diameter in
Beauveria spp. to up to 200 μm in Tuber (ascospores), Mucorales (zygospores) and Endogone
15
(chlamydospores) (Weber & Hess 1976). Among the 303 European polypores investigated by
Kauserud et al. (2008) 85 % of the species have spores with a volume less than 100 μm3.
Chaetoporellus latitans has the smallest spores (0.94 μm3), while Perenniporia ochroleuca has
the largest spores (548 μm3). These volumes (100, 0.94 and 548 μm
3) correspond to diameters of
approximately 6 μm, 1 μm and 10 μm of spherical spores. Species with larger basidiocarps tend
to produce larger spores; Parasitic species have larger spores than saprotrophic species; Species
on deciduous wood have larger spores than species on coniferous wood; White rot species have
larger spores than brown rot species on the same kind of wood (deciduous and
coniferous)(Kauserud et al. 2008). Basidiospores in the species of hymenomycetes (including
e.g. the orders Agaricales, Russales and Boletales) are rather small (mean length = 8.4, width =
4.9 μm and volume = 156 μm3) and elongated (mean L/W quota = 1.8) (Parmasto & Parmasto
1992).
Diaspore longevity
To disperse in space as well as in time it is necessary that spores can survive the conditions in the
air and in the ground. The spore may encounter drought, UV-light and freezing in the
atmosphere. Zanten (1977; 1978) tested the viability of spores from several species from the
southern hemisphere and New Zealand as well as from the northern hemisphere after drought,
dry freezing and wet freezing up to three years. He concluded that the species with restricted
distributions stayed viable for a shorter time than those that occurred on different continents. He
also concluded that the species from the northern hemisphere on average stayed viable for a
longer time. Malta (1922) tried to germinate moss spores from herbarium specimen of different
age. Most collections that were older than 10 years did not germinate, and most of the spores that
germinated were younger than 5 years. The oldest collection which contained viable spores was
16 years (Ceratodon purpureus). This result contradicts the fact that weedy plants like
Ceratodon purpureus and Funaria hygrometrica are seldom found in the diaspore bank. During
(1987) tried to investigate this by sowing spores of Funaria hygrometrica in a chalk grassland.
The spores germinated in the greenhouse still after two years but not in the field. Löbel (2009)
noted a 30–40 percentage decline in viability of spores during 50 days of dry storage.
16
Whitehouse (1984) found that tubers of Dicranella staphylina were still viable after 50 years of
storage. Sundberg & Rydin (2000) showed that 15-35 % of Sphagnum spores survive 13 years of
storage under humid conditions.
Spores and fragments were stored in water for up to six months of the following five mosses:
Schistidium rivulare, Racomitrium aciculare, Dicranoweisia crispula, Oligotrichum hercynicum
and Ceratodon purpureus by Dalen & Söderström (1999). The spores survived longer when
stored under dry conditions compared with storage in water except for in Schistidium rivulare,
whereas the fragments did not have a higher survival stored under dry conditions (Dalen &
Söderström 1999).
The spores of the fungus Beauveria bassiana showed 90% germinability after 635 days in
darkness at 0% relative humidity and 8 °C. However within 28 days in light at 75 % relative
humidity at 18°C none were germinable. (Clerk & Madelin 1965)
Number of diaspores
The number of spores per capsule is substantial. Ingold (1974) states that the capsule of
Eurhynchium confertum contains between 250,000 and 750,000 spores. Kreulen (1972)
estimated the number of spores in one capsule to be 450,000 in Atrichum undulatum and 1.4
million in Polytrichum juniperum. Buxbaumia viridis has 6 (1.4–9) million spores per capsule
(Wiklund 2002). Sundberg & Rydin (1998) counted the number of spores in eight peatmosses
and estimated from 18,500 spores per capsule in Sphagnum tenellum to 240,000 in Sphagnum
squarrosum. For additional estimations of the number of spores per capsule see Fig. 3. More
interesting for dispersal is the spore output per area unit. Sundberg (2002) studied several
Sphagnum species and reported an annual production of 16 million spores per square metre of
mire. Longton (1976) estimated the spore production of Pleurozium schreberi at one site to be
100 million spores per square metre. The spore production of the liverwort Ptilidium
pulcherrimum varied from approximately 20 to 300 million spores per square metre colony
during a four year period and the production per square metre of forest land was estimated to
75,000-150,000 (Jonsson & Söderström 1988). Spore production estimations have been done for
17
three moss species in millions per square metre of substrate: 37 for Atrichum undulatum, 7,400
for Grimmia pulvinata and 38,300 for Tortula muralis (Longton & Schuster 1983). So even if
the vast majority of the spores are deposited very near the mother colony there are many that can
contribute to the LDD.
Figure 3. The number of spores per capsule (y-axis on a logarithmic scale) versus the mean spore
diameter (x-axis in μm). The data have been compiled from several sources (Ingold 1959; Schuster 1966;
Kreulen 1972; Ingold 1974; Longton 1976; Mogensen 1978; Söderström & Jonsson 1989; Miles &
Longton 1992; Boros et al. 1993; Sundberg & Rydin 1998; Wiklund 2002; He & Zhu 2010; Cuming
2011) and are based on estimates from 92 species with many species from some genera e.g. Riccia and
Sphagnum.
Diaspore banks
Bryophytes may survive in the soil as fragments of e.g. the stem, spores and specialized asexual
propagules such as tubers or gemmae. It may facilitate species' survival under unfavourable
conditions. This makes it more difficult to interpret and investigate dispersal in space under
certain circumstances.
18
Dominant, common pleurocarpous mosses seem to be rare in the diaspore bank, while short-
lived species that only are present above ground some time after disturbance seem to be more
common in the diaspore bank. A higher proportion of larger than of smaller diaspores is
represented in the diaspore bank of bryophytes. The opposite is true for vascular plants. As
mentioned above not only spores can constitute the diaspore bank. For example Barbiolphozia
lycopodioides was present in the diaspore bank in forests in northern Sweden even if it does not
produce gemmae and sporulates very rarely (Jonsson 1993). Most studies of diaspore banks
have consisted of cultivating soil samples (During 1997; During 2001). To assess if the diaspore
bank consists of spores or vegetative parts of the bryophyte is therefore difficult. It has been
proven that Sphagnum can regenerate from the diaspore bank Clymo & Duckett 1986; Sundberg
& Rydin 2000). The latter buried spores at different depth and that the survival was best under
wet but aerobic conditions and that the spores died within 2-3 years under anaerobic conditions.
Diaspore banks can be classified as transient, short-term persistent or long-term persistent
(During 2001) (Table 2). Since a direct measurement of the diaspore age is difficult to acquire,
the age must be assessed by means of indirect estimates such as the time since the species was
observed last or some approximation of humus accumulation rates.
Table 2. Different types of diaspore banks with some examples among the bryophytes.
Diaspore banks Time frame Example species Spore size
transient months
short-term persistent 1–5 years Anthoceros Anthoceros punctatus 36–60
μm Phaeoceros (30–)38–56(–
75) (Zanten & Pócs 1981)
long-term persistent >5 years Physcomitrium
sphaericum
–30 μm
19
Frahm (2006) suggested that the occurrence of several rare thermophilic openland species
(Microbryum curvicolle, Acaulon triquetrum, Weissia longifolia) on the edge of a forest road
could be interpreted as coming from a diaspore bank from the time when a wineyard had been
situated at the site almost 100 year old ago. The species are very rare in the area and the nearest
known recent site is 40–100 km away.
Dormancy
Most bryophyte diaspores seem to have enforced dormancy, i.e. they start to germinate when
suitable conditions concerning e.g. light and water are available. There seems to be some
exceptions. The disc-like gemmae in Blasia pusilla do not seem not able to germinate right away
after dispersal (During 2001). Laaka-Lindberg & Heino (2001) studied the gemmae of Lophozia
silvicola and suggested that part of them enter dormancy. Sundberg & Rydin (2000) noted a
better germination of 3 years old spores than 2 years old ones and concluded that it could be due
to the wet and cool conditions during the third year and the spores' need to be hydrated for a long
period and possibly chilled to germinate.
Deposition
Deposition of small diaspores is highly affected by the weather conditions. Turbulent diffusion
and precipitation are two important factors (Schmidt 1967). A small object collects diaspores
better than a large object due to the thickness of the boundary layers around objects. With
increasing wind velocity the boundary layer becomes thinner. A beech pollen with a diameter of
42 μm settles more easily than a birch pollen with a diameter of 21 μm but the difference
decreases with increasing wind velocity (Whitehead 1983).
Besides the source strength, the amount of suitable target habitat in an area is determining the
probability of realized dispersal because the number of diaspores that reaches a suitable habitat
varies with the area of that habitat if we assume a random dispersal.
n/N~a/A
20
where N=the total number of diaspores, n=the number of effective diaspores, a=the area of target
surfaces/suitable area, A=the total area of the surroundings (vegetation and habitat). a (the total
area of stigmata) is lower for pollination than for most other kinds of spore dispersal where the
total area of the target substrate often is larger. (Faegri & Pijl 1979)
How to study dispersal
Dispersal can be studied by direct or indirect methods. The direct methods can be divided into
Lagrangian methods and Eulerian methods. The first method include tracking a single diaspore
and the second trapping diaspores
Direct methods
Lagrangian methods – tracking individuals/single diaspores
These methods include measuring how far, how fast and in what direction individuals move and
have the advantage that it is easier to track LDD events. This kind of approach is used in studies
first and foremost on large animals with radio or satellite tracking, but experiments with insects
has also been performed (Nathan et al. 2003). It is also easier to detect clumped patterns in the
dispersal of diaspores with these kinds of methods. It may also be applied to plants dispersed by
animals and humans. (Bullock et al. 2006)
The movements by stolons of clonal plants such as Trifolium repens can be backtracked by
looking at the annual growth of the individual (Cain et al. 1995), but this will be restricted to
short distances and cannot be defined as LDD by any means.
As Langrangian methods are rather labourious the number of diaspores that can be followed is
limited (Nathan et al. 2003). Larger diaspores, like those with pappi in e.g. Cirsium arvense and
Senecio nemorensis, can be followed individually (Tackenberg 2003). However as the size of
the diaspores decreases it becomes more difficult to follow them. Hence no studies using these
kinds of methods have been done on small wind-dispersed diaspores.
Release of marked diaspores is an approach that is somewhere in between the Langrangian and
the Eulerian approach. But as diaspores rarely are marked individually and the trajectory from
21
release to catch is not observed these methods should be categorised as Eulerian methods.
Studies of insect dispersal of pollen have used coloured powder. Other studies have used
artificial propagules to mimic dispersal of e.g. seeds in rivers (Andersson et al. 2000).
Eulerian methods – trapping
Here, the information of the diaspores' whereabouts between release and capture is lacking.
Since only a small proportion of the surface is sampled, the diaspore source needs to be very
strong to detect LDD (Nathan et al. 2003).
Many authors have studied the deposition of spores around a central colony using microscope
slides with some sticky substance such as melted petroleum jelly. Stoneburner et al. (1992)
studied the deposited spores of two colonies of the acrocarpous moss Atrichum angustatum up to
15 m. The traps at 1.5 m right above the colony did not catch any spores but those at the other
distances (0.5, 1.5, 2.5 and 15 metres) caught spores at both 0.5, 1 and 1.5 metres height.
Söderström & Jonsson (1989) studied the spore deposition around one colony of the leafy
liverwort Ptilidium pulcherrimum in 7 distance intervals at 49 collection points up to 10 m from
one colony. They concluded that 43 % of the spores were deposited within 2.5 m from the
colony. Sundberg (2005) studied the dispersal of six Sphagnum species, which have an active
release, up to 3.2 m from a central colony and found that 8 to 32 % of the spores which were
dispersed outside the colony did not travel farther than 3.2 m. Miles & Longton (1990) trapped
spores from the acrocarpous mosses Atrichum undulatum and Bryum argenteum up to 2 m from
a colony and estimated that 7.2 % to 12.7 % and 2.2 to 4.5 % of the dispersed spores were
deposited within 2 m from the colony. In contrast 70 % of the spores were estimated to be
deposited within 2 m of a colony of 46 sporophytes of the moss Tortula truncata (Roads &
Longton 2003). Pohjamo et al. (2006) compared the dispersal of spores of equal size and
gemmae (diameter = 9–12 μm) of the liverwort Anastrophyllum hellerianum. They concluded
that between 17.5 and 43.1 % of the spores released each day and between 0.64 and 4.8 % of the
gemmae available were deposited within 10 m from the central colonies. The percentages for
spores were calculated of the released spores, whereas the percentages for gemmae were
calculated of the total of available gemmae and included also gemmae that remained attached to
the shoots after the study period. Since the ratios were calculated in different ways they cannot
22
be compared directly. Rain did not affect the dispersal pattern significantly. More gemmae were
however released during rainy days.
The problem with many small wind-dispersed diaspores is that they are difficult to recognize and
determine to species level. One way is to remove all possible spore sources within a certain
radius or study realized dispersal where the spores have germinated and become easier to
identify. Kallio (1970) used petridishes with agar and wood discs up to 200 m from a spore
source to study the spore dispersal of the root-rot fungus Heterobasidion annosum. He exposed
agar plates up to 2 minutes and the wood discs up to 2 hours. Under the colony and at 1 m the
petridishes were exposed for 5-10 seconds. The deposition were here estimated to at least
200,000 spores/dm2/hour and 30,000 spores/dm
2/hour. The deposition at 50, 100 and 200 m were
similar and varied from 0 to 49 spores/dm2/hour calculated from an exposure time of 1-2
minutes. Mycologists can also study a sort of realized dispersal using traps with monokaryotic
mycelia and check if they have been dikaryonized by looking for clamp connections or other
morphological changes in the mycelia after an incubation time in a laboratory. There strains are
selfincompatible, so the arriving spore must be of a different mating type in order for a
dikaryotic mycelium to develop (Edman & Gustafsson 2003). The monokaryotic mycelia have
been cultivated on nutrient agar (Adams et al. 1984; Williams et al. 1984) but Edman &
Gustafsson (2003) cultivated monokaryotic mycelia of some polypores on wood-discs instead.
Norden & Larsson (2000) studied dispersal of the wood-inhabiting polypore Phlebia centrifuga
using single spore mycelia on agar and obtained dicaryonized mycelia up to 100 m using ten fruit
bodies as a source in a circle where all the fruitbodies within 200 m from the center had been
removed. In a second experiment they trapped spores at 0–350 m and at 500–800 m from the
limit of a nature reserve with old-growth forest with a more diffuse spore source. In this
experiment they detected spore deposition on 15 of 60 in the first distance and on 4 of 59 in the
second distance.
Hallenberg & Kuffer (2001) also used petri dishes with single spore mycelium to trap spores
from eight wood-inhabiting basidiomycetes. They were exposed on a roof in Gothenburg for 24
h six times during October and November. They detected spores from those species that were
23
common in the vicinity as well as the species that had the nearest known site 1000 km to the
south as well as to the east.
One possibility to be sure that the trapped diaspores originate from the mother colony is to apply
15N isotopes to the mother plants during the flowering stage so the seed will also have higher
content of this isotope than other plants. Carlo et al. (2009) applied three urea doses to plants of
Solanum americanum and Capsicum annuum and concluded that it was possible to detect this
enrichment even in seedlings. Since many spores are very small compared to the stages after
germination and that at least many bryophytes live in nutrient poor environment it still remains
to be tested if this method is applicable for studies on spore dispersal.
Another method for tracing the origin of diaspores is to use genetic markers. By analysing the
pericarp, which is inherited maternally, it is possible to determine from which of the sampled
mother plants the diaspore originates or if it has dispersed from some unsampled plant outside
the study area. This is feasible for large seeds like acorns that Grivet et al. (2005) studied with
microsatellites and found that the dispersal distance was not more than 100 m. However, for very
small seeds and spores this method is not applicable. Instead one could sample all possible
parents, both males and females, as well as seeds and/or seedlings in an area or population. As
this will lead to extensive sampling only small areas/populations can be investigated with this
approach. The "Two-Gener" method includes collecting seeds from the mother plant and by
analysing pollen genetic structure estimate the average pollen dispersal distance by sampling
possible father plants in the vicinity (Broquet & Petit 2009). Parentage analysis has also been
used to study pollen dispersal in a similar way. Oddou-Muratorio et al. (2010) used
microsatellites to investigate dispersal of seeds and pollen in Fagus sylvatica and F. crenata by
estimating the historical and recent gene flow.
Indirect methods
Genetic similarity between populations can be used as a measure of the connectivity between
populations and assess historic dispersal. The easiest way is if the different genotypes could be
distinguished with the naked eye. An elegant way to study realized pollen dispersal was used by
24
Bannert & Stamp (2007) who used differently coloured maize to detect pollen dispersal between
fields. In a valley in Switzerland they planted one field with yellow maize and at several
distances (50–4500 m) from this field they planted white maize. Cross-pollination, which could
be detected easily by yellow kernels, was detected at 50–371 m in the main wind direction.
However, one is rarely fortunate enough to have such traits available. Neutral genetic markers
are an increasingly used tool in the study of dispersal in recent decades. The price, the amount of
information generated, the ability to detect heterozygotes (codominance) and the specificity (and
hence the risk of contamination) are some factors to take into account when choosing the
method.
Enzymes with the same function can be separated on gels with an electric current due to their
different size and charges. Isozymes are enzymes with the same function coded by different loci
whereas allozymes are variant coded by different alleles in the same loci. Isozymes/allozymes is
a rather cheap method where you separate different variations of an enzyme (allozymes) on a gel.
The intercontinental genetic structure of the basidiomycete Schizophyllum commune was
investigated by James et al. (1999) using sample strains from North America, Central America as
well as from South America, Asia, Africa, Europe and Australasia. Since there was a higher
genetic variation within than between intercontinental populations they concluded that LDD
could be efficient enough in this species to secure a gene flow even over large distances.
Grundmann et al. (2007) studied the moss Pleurochate squarrosa in the Mediterranean area
using both allozymes markers and DNA sequence data. The data from allozymes showed no
isolation by distance but the sequence data did.
Thirty-two populations from Asia, America and Europe of the copper-moss Scopelia cataractae
were investigated for 15 allozymes loci. It was concluded that the species was native to America
and Asia and that LDD has been important for the species. (Shaw 1995)
Different types of PCR-based fingerprinting methods have become more used recently. They can
generate more information than alloenzymes. Some are not codominant like RAPD (Random
Amplified Polymorphic DNA), AFLP (Amplified Fragment Length Polymorphism). RFLP
25
(Restriction Fragment Length Polymorphism) is often codominant. Restriction enzymes cut the
DNA at certain sequences. The fragments of different lengths are separated on a gel.
Microsatellites (ISSR – Inter Simple Sequence Repeat) are codominant. In opposition to the
methods mentioned above microsatellites can reveal the relationship to other alleles (Silvertown
& Charlesworth 2001). SNP (Single Nucleotide Polymorphism) is a novel method that has not
been used in so many studies on bryophytes yet.
James & Vilgalys (2001) trapped spores of the common basidiomycete Schizophyllum commune
using haploid mycelia on agar in petri dishes exposed for 12 hours in five out of seven sites from
northern South America through the Caribbean to southeastern USA and calculated from this a
deposition of 18 spores/m2/hour. The samples were taken in areas where no fruit bodies were
expected to occur in the closest vicinity (non-forested areas) so the result should describe the
background deposition. They analysed five loci of the dikaryotic mycelia with RFLP. Dispersal
over the Caribbean Sea could not be showed unambiguously. However, they showed in one locus
a possible pattern that could be interpreted such that gene flow had occurred from the south to
the north (which was the prevailing wind direction) but not the other way around.
Snäll et al. (2004) studied the genetic structure of Orthotrichum speciosum and O. obtusifolium
in a forest landscape in Finland using AFLP. They concluded that both species were limited by
dispersal beyond 300-350 m.
Szövényi et al. (2008) investigated the genetic structure for six peatmosses with amphi-Atlantic
distribution using microsatellites and concluded that both migration and the polymorphism of the
founders can explain the genetic structure of the intercontinental populations. The moss
Polytrichastrum formosum showed a low genetic differention ( showed both with allozymes and
microsatellites) between three populations in Denmark and three in the Netherlands
approximately 450 km apart and Van der Velde et al. (2001) concluded that this was due to spore
dispersal.
Other genetic methods can be used to study dispersal over longer time spans. For phylogenetic
studies and immigration history of a species in different areas, sequencing of certain parts of the
genome has been used, e.g. ITS (Internal Transcribed Spacer) a non-coding RNA part of the
26
Ribosome DNA. Huttunen et al. (2008) studied Homalothecium and used a substitution rate of
0.8–2.0 %/106 year and a mean of 0.014 substitutions/site
/10
6 year based on (Bakker et al. 1995)
for green algae and higher plants. Moncalvo & Buchanan (2008) studied the basidomycete
Ganoderma applanatum-australe species complex and refers to a substitution rate of fungi of
1.26 % /108 year and concludes that there is a occasional gene flow due to LDD in the southern
hemisphere after the separation of Godwanaland in this species.
Biogeographical studies
Dispersal and vicariance are two factors that have been used to explain species distributions. The
number of endemic taxa is lower in ferns (34%) compared with angiosperms (80 %), which can
be explained by: 1. that many fern taxa are older. 2. speciation rate is lower for ferns 3. ferns are
better dispersers (Smith 1972 in Wolf et al. 2001). This can be said of many other organism
groups with small diaspores. Factors that could be contributing to the opinion that all micro-
organisms are omnipresent, are that distribution data is incomplete for many species and that
many rare species are still to be described. Some flagship species have restricted distribution, and
even if many species have drought resistant diaspores, such as cysts, others have more sensitive
diaspores. The fraction of endemic micro-organisms has been assumed to be 30 %. (Foissner
2006)
A comparison of the composition of floras among continents or islands can be used as an indirect
way of detecting dispersal and connectivity between land areas. For example, Munoz et al.
(2004) showed a close correlation between the predominant wind direction in the summer and
the composition of bryophyte flora in the southern hemisphere. Zanten & Pócs (1981) compared
the composition of bryophytes of the southern hemisphere. The sensitivity of the spores to
drought and freezing was investigated in several species. They concluded that the endemic or
geographically restricted species had spores with lower longevity than those with a wider
distribution. Species that are endemic but have long-lived spores may have been overlooked and
may in fact occur in other continents than hitherto known.
27
Other studies
Smaller scale patterns can be used for interpreting shorter time scale processes. Miller &
McDaniel (2004) studied the moss flora on calcareous mortar along a highway in a landscape
with only acidic substrata. During the 65 years since the construction of the highway a number of
calcicolous bryophytes that did not occur in the vicinity earlier had colonised the calcareous
substrate and many species had dispersed 5 km or more during this time span.
Hutsemekers et al. (2008) compared the bryophyte species on slag heaps of different ages (aged
by dendrochronology of rooted trees) with the nearest 4x4 km grid cell with occurrence of the
species according to a bryophyte atlas of Belgium. They concluded that the area of the patch and
time available for colonization were important for the number of species and that 56 % of the
species emanated from a source located 6-86 km during the last 50 years.
The two mosses Campylopus introflexus and Orthodontium lineare were accidently introduced
to Europe from the southern hemisphere. By documenting the first finding of the species in
provinces across Europe it is possible to get a approximate estimation of how fast the species
disperse across the continent (Hassel & Söderström 2005).
Lichen species were inventoried on 94 gravestones in five counties in England, and classified
according to which 20 year period they were raised (1650 to 1980). Abundance (=average ratio
of gravestones with the species for all periods that it occurred), area of occupancy (=number of
squares in Great Britain), spore size and colonization ability (=e.g. the most recent period with an
occupied stone) were noted for 92 saxicolous species. Area of occupancy was positively
correlated with both abundance and colonization ability, while spore size was not correlated to
any of the parameters. (Leger & Forister 2009)
By studying the distribution of trees and the occurrences of the two epiphytic mosses
Orthotrichum speciosum and Orthotrichum obtusifolium in a forest area, Snäll et al. (2003)
found that the occurrences of the spore dispersed O. speciosum was influenced by connectivity,
while the gemmiferous O. obtusifolium was not. They also predicted that 17 % of the
colonizations of O. speciosum occurred farther away than 15 m (Snäll et al. 2003). Hedenås et al.
(2003) noted that the sexually dispersed epiphytes Collema curtisporum and Orthotrichum
28
speciosum were more aggregated than their host trees Populus tremula while three asexually
dispersed species C. furfuraceum, Leptogium saturninum and O.obtusifolium were not.
Models
Phenomenological models try to fit the best model disregarding other measurements than
distance, while mechanistic models try to find the best model from measured parameters like
wind speed and abscission height. There are also intermediate models which e.g. Snäll et al.
(2007) have proposed.
When comparing functions from different studies it is important to be aware of that several
different functions are used. In response to the confused use of dispersal curves and dispersal
kernels, Cousens et al. (2008) used probability density function (pdf) as a general term and
recognized three functions (distance pdf: f(r), density vs distance relationship: ωθ(r) and density
pdf: g(r). A function of one type of distribution should not be compared to any of the others
unless they have been transformed. The function that describes the probability of single diaspore
to arrive at a certain distance is called the "distance pdf". Taking samples in form of e.g. traps
will generate a density vs distance relationship. The probability density function for a diaspore
landing in an infinitely small area at distance r from the origin is called the "density pdf". Power,
negative exponential, Weibull, half normal, half student t and Cauchy have been used to describe
density vs distance relationship. A possibility, especially to describe fat-tailed curves, is to use a
mixed model, i.e. combining two functions. (Cousens et al. 2008)
Phenomenological models
Fitt et al. (1987) compared 325 data sets with deposition of pollen, micro-organisms and spores.
Power law and exponential models were fitted to these. The fit was similar for both models in
most cases. However, fungal spores (<10 μm) fitted better with the power law function and
droplets with exponential. A curve should not be extrapolated beyond the available field data.
Bullock & Clarke (2000) fitted negative exponential and inverse power models to a data set of
seed dispersal of Calluna vulgaris and Erica cinerea up to 80 m from the source. Even if the
inverse power fitted the data better none of the models estimated the tail very well. A mixed
model was used to get a better fit in the entire span of the function.
29
Mechanistic models
Different functions and parameters have been used. Dispersal models often include properties of
the diaspore such as terminal velocity, as well as meteorological and landscape properties such
as wind speed. Tackenberg (2003) found that the thermal updrafts and other turbulence had a
large influence on the rate of LDD. The question is what is most important in LDD: extreme
windy conditions during hurricanes and tempests or warm conditions with convection. Horn et
al. (2001) also emphasised the effect of updrafts on winged seeds and even if some diaspores of a
certain species fall faster than others every diaspore can be caught in an updraft and contribute to
LDD.
One important factor is the weather conditions. The laminar layer during daytime in
roughweather is less than a millimeter whereas it during calm weather at night is much thicker.
Spores released at midday have a better chance of being transported far away. Conditions of low
humidity tend to be correlated with turbulence. (Ingold 1974)
The Gaussian plume model (GPM) has been used for estimating concentrations of gases and
particles up to 10 km from a point source. Spijkerboer et al. (2002) calibrated the model with the
weighted least-squares method. They estimated the escape fraction outside the canopy to 64 +-
17% from a experiment where spores of Lycopodium clavatum were released and trapped with
35 samplers up to 100 m in a potato field. Katul et al. (2005) used a mechanistic analytical model
which has an asymptotic power law tail with an exponent of -3/2 and tested it on datasets on
seeds from trees, shrubs and herbs.
Conclusions
Many species with small diaspores produce a large number of diaspores which can constitute a
strong source for LDD even if only a small proportion of the diaspores is transported far away
from the mother plant. The studies that have been performed give somewhat contradictory
answers how effective these species are as long-distance dispersers. On which temporal and
spatial scale that species are dispersal limited needs further investigation.
30
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