Plant Ecol (2007) 192:45–53

DOI 10.1007/s11258-006-9225-1

ORIGINAL PAPER

Impact of ploughing on soil seed bank dynamics

in temporary pools
Vincent Devictor Æ Jacques Moret Æ
Nathalie Machon

Received: 31 May 2005 / Accepted: 23 October 2006 / Published online: 16 November 2006

Springer Science+Business Media B.V. 2006

Abstract We examined the impact of ploughing unploughed pools. Our results agree with the
on soil seed banks of plant communities living in temporal storage effect generally suggested to
temporary marshes located in agricultural fields. describe the seed bank property of plant commu-
The quantity, quality and vertical distribution of nities. But in addition, we showed that ploughing
seeds were quantified under ploughed or un- induces a spatial storage effect in accumulating
ploughed treatment at community level. We also species and individuals in the seed banks that
focussed on a typical semi-aquatic species, favourably influence community dynamics. We
Damasonium alisma, to investigate the impact conclude that, in contrast to what is usually
of ploughing at population level. We used two thought, ploughing disturbance can be of benefit
complementary techniques to study seed banks: for such ephemeral wetland vegetation.
hand sorting and seedling emergence. We found
that species richness of seeds, seed abundance Keywords Agricultural practices
Community

and germination ability were strongly affected by Damasonium alisma
Ephemeral vegetation

ploughing at community level. Concerning Storage effect
Wetland
D. alisma, most of the seeds (56%) were stored
in the two deepest soil layers among the four
considered in ploughed pools. Moreover, the Introduction
germination rate was higher for buried seeds
(84%) than for seeds collected at the surface Several species have evolved with human activ-
(33.6%). These patterns were almost inverted in ities, and many of them can even be threatened
by the abandonment of anthropogenic practices.
V. Devictor (&)
N. Machon This link has been primarily studied in agricul-
UMR 5173 CNRS MNHN Conservation des Espèces, tural landscapes, where biodiversity and human
Restauration et Suivi des Populations, Muséum practices are strongly related (Stoate et al.
National d’Histoire Naturelle, 61 Rue Buffon,
2001). Through their impact on population
75005 Paris, France
e-mail: [email protected] dynamics, agricultural practices have contrib-
uted to shape species richness and distribution
V. Devictor
J. Moret
N. Machon of uncultivated plants. Temporary pools located
UMS 2699 Inventaire et Suivi de la Biodiversité,
in arable fields provide a good illustration of
Conservatoire botanique National du Bassin parisien,
Muséum National d’Histoire Naturelle, this relationship between agricultural practices
55 Rue Buffon, 75005 Paris, France and plant communities. Their drainage was

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recently undertaken, inducing the decline of kill weed seeds and seedlings before cultivated
such pools (Gallego-Fernández et al. 1999; Le- plants are sowed (Roberts 1981; Beuret 1989).
feuvre et al. 2000). Consequently, the flora However, the consequences of this agricultural
adapted to this type of environment, composed practice on community dynamics in wetland
mainly of small annual plants, is considered to communities are not clearly established. For
be threatened, and many of such species are wetland species affected by ploughing, a seed
legally protected. bank is likely to act as an effective source of
Among agricultural practices affecting plant colonising species and to determine the above-
communities, ploughing is one of the most ground species composition even after deep
deleterious since it physically removes seeds ploughing (Luzuriaga et al. 2005).
and seedlings from the soil surface (Ghersa and To study seed banks, two techniques have
Martı́nez-Ghersa 2000; Tørresen and Skuterud been commonly used: hand sorting and seedling
2002). On the other hand, seeds play a key role emergence (Standifer 1980). Usually, each of
on individual’s and species’ survival (Shaukat these two techniques appears to be insufficient
and Siddiqui 2004; Adams et al. 2005). Soil seed by itself. Seedling emergence gives information
bank was therefore usually seen as a basic way about the plant community; it reveals a large
to momentarily escape unfavourable environ- part of the species contained in the soil samples
mental conditions as severe drought or frost and their interactions during germination are
(Cohen 1966). More recently, Stöcklin and Fish- more or less preserved. However, many seeds
er (1999) showed that species with high seed cannot germinate in ex situ conditions, and
longevity had lower extinction rates. On an some of the species are not detected with this
evolutionary scale, seed bank and dormancy technique. Consequently, seed hand sorting is
were also presented as a way to face unpredict- often more accurate. Unfortunately, this meth-
able environmental changes (Pacala 1986; Thrall od of direct counting gives no information on
et al. 1989), to maintain genetic polymorphism the ability of seeds to grow in their natural
or to promote species diversity (Templeton and conditions, and it breaks the natural vertical
Levin 1979; Hedrick 1995; Vitalis et al. 2004). structure of the seed bank. Furthermore, very
These considerations have been mainly theoret- small seed species can remain undetected or
ical and rarely tested in natural populations unidentified with this technique. The cultivation
through experimental approaches (but see Kalisz of the seeds is thus often necessary afterwards.
1991; Bliss and Zedler 1998; Bonis 1998). At the In fact, these two techniques are complemen-
community level, the positive effects of seed tary although in most cases they are not utilised
bank properties have also been used as a together.
relevant tool in conservation and restoration The aim of this paper is to understand the
programmes (Miller and Cummins 2003) or impact of ploughing on seed banks of tempo-
ecosystem management (Wienhold and van der rary pools located in arable fields at community
Valk 1989; Warr et al. 1993; Jalili et al. 2003; and population levels using both seedling
Middleton 2003). However, none of these previ- emergence and soil sieving. An experimental
ous studies was done on wetland communities approach was used to compare the quantity,
affected by ploughing. the vertical distribution and the quality of seeds
In natural wetland communities, seedling between ploughed and unploughed areas. We
emergence is usually shown to be strongly related addressed two specific objectives: (1) What is
to the fluctuating water level (van der Valk 1978). the impact of ploughing on the seed banks’
But in cultivated fields, the seed bank dynamics viability for plant communities living in tempo-
may completely differ from what was found in rary pools? (2) What is the impact of ploughing
uncultivated lands since ploughing may have a on a typical ephemeral threatened wetland
deep impact on seed distribution and viability. species, Damasonium alisma, at the population
Indeed, the main role of ploughing is precisely to level?

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Methods decreased during the twentieth century because

of the disappearance of wet zones in Europe
Pool description (Wheeler 2000). This semi-aquatic herb is found
in England, France, Italy, Spain and possibly
Clay layers induce the formation of temporary south-western Asia (Birkinshaw 1994). It is
pools in arable fields. Such pools generally fill legally protected in France (Danton and Baffray
up during the winter period (for a 30 cm 1995).
maximum depth during the study) and progres-
sively dry up during spring. Two of these pools Seed bank study
located in the agricultural neighborhoods of
Paris (France) were studied: one near the city Our sampling method was strongly influenced by
of Echarcon (E) and the other near the city of the fact that we worked on protected species
Fleury-Merogis (F). Both pools E and F were (seed samples had to be few). However, the
round and had approximately the same size of small size of the pools allowed us to get
50 m2 in March 2003. The E pool was located sufficient information with few samples. Ten
exactly at the edge of a cultivated field and replicate soil samples (cylinders of 7-cm-diame-
consequently one half was located in the culti- ter and 15-cm-depth) were cored in each pool.
vated area (called Ec for E cultivated) and the This sampling was performed in March 2003, i.e.
other half was located in an uncultivated area after fall ploughing and before 2003 seed pro-
(called Ew for E wild area). This pool was duction. Each sample was then divided into 5
located in a typical intensive farmland landscape depth layers of 3-cm-thickness each, from the
of annual arable crops. The land-use past of this top to the bottom (0–3, 3–6, 6–9, 9–12, and
pool remained similar for a period longer than 12–15 cm). Such division was limited in un-
5 years, but the annual species grown in the ploughed pools (F and Ew), in which the soil
cultivated area may have changed each year. was so compact that the deepest layer
The F pool was located in a wild area, which (12–15 cm) was hardly reached and therefore
had not been cultivated for at least 5 years. The not considered. Finally, the soil of each layer
F pool was situated 5 km away from the E pool. was sieved through 500- and 250-lm-aperture
The three pools had the same type of soils meshes. All seeds of each species collected by
(uniformity of edaphic conditions) and provided this technique were hand sorted and identified.
our natural experimental framework to test the One hundred and fifty D. alisma seeds were
influence of ploughing on temporary wetland isolated from each layer and cultivated. Culti-
communities. vation was performed in waterproof pots placed
in incubators (12 h of light at 15C and 12 h of
Species description obscurity at 10C), filled with sterilized pool
soil. Since D. alsima seeds were shown to
All ephemeral semi-aquatic species growing in germinate only below water and not just in
temporary pools were studied. Among all spe- damp conditions (Birkinshaw 1994), each pot
cies, Damasonium alisma Miller (Alismataceae), was identically watered (i.e. the soil was kept
the star fruit, was more specifically considered under 1 cm of distilled water, pH = 7), and
for some experiments. This species is a rare randomly positioned. Seedlings were numbered
annual plant growing in the muddy margins of after 40 days.
pools with seasonal fluctuating water levels. It is To study seedling dynamics for the whole
a small (5–30 cm high) white flowered plant. community, 15 other soil cores (cylinders of
Seeds germinate in early spring only under 20-cm-depth and 5-cm-diameter) were randomly
water, and the plant finishes its cycle before extracted from each part of the E pool (Ec and
the pools have completely dried during summer. Ew) and from the F pool in March 2003. Each
Its mating system is presumably facultative sample was then divided in two layers: an upper
autogamous (Vuille 1987). Its distribution has stratum (U) and a lower stratum (L) of 10-cm-

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thickness each (0–10 and 10–20 cm). After adjust- Table 1 Effects of pool, depth and interaction between
ment of their weight, each layer was spread out in pool and depth on the number of seeds found in the core
samples by hand sorting (log transformed)
large waterproof plastic pots and placed in the
garden of the National Botanical Conservatory of Source DF F Prob > F
the Parisian Basin. Each layer was watered
Pool 2 2.67 0.0705
regularly to keep the soil surface under 1 cm of Depth 1 93.63 <0.001
water. The location of each pot in the garden was Pool · depth 2 46.00 <0.001
regularly randomised. Pots with sterile soil were Pool and depth were considered as fixed effects and core
also cultivated to check for seed contamination as a random effect of the mixed model (LME S-PLUS)
from the garden. Emergence of seedlings was
assessed weekly during 6 weeks. Results

Data analysis Impact of ploughing on wetland community

First, we wanted to test the effect of ploughing on The number of seeds was not markedly different
seeds distribution along the vertical. The number among pools (P = 0.07) but depended strongly on
of seeds collected from each depth layer obtained the depth at which they had been cored
from Ec, Ew and F after hand sorting, was (P < 0.001) (Table 1). Moreover, the effect of
compared in each pool and between pools. Since depth was contrasted among pools (interaction
each depth layer was not independent from one Pool · Depth, P < 0.001). In particular, more
another but was grouped according to the core seeds were found in the upper soil layers (i.e.
sample, we performed a linear mixed model. near the surface) in the fallow parts of the pools,
Mixed-effect models extend linear models by whereas in the cultivated parts of the pools, more
incorporating random effect to account for cor- seeds were found in lower layers (Fig. 1).
relation among observations within the same
group (Pinhero and Douglas 2000). Therefore, Seedling emergence
we incorporated ‘‘Depth’’, ‘‘Pool’’, and interac-
tion between ‘‘Pool’’ and ‘‘Depth’’ as fixed effects No germination occurred in the sterilised soil
and ‘‘Core’’ as a random effect. We used the same pots. In the E pool, all the species that emerged
method with number of seeds of D. alisma only. from the cores extracted from the cultivated pool
The number of seeds was log transformed for (Ec) were also present in the cores coming from
these analyses. To test the effect of ploughing on Ew: Lythrum hyssopifolia L., Lythrum portula L.,
D. alisma seed quality, we performed a regression Juncus tenageia L., Polygonum persicaria L.,
analysis on the germination percentage of the 150 Gnaphalium uliginosum L., Damasonium alisma
seeds we had cultivated, according to depth Mill. (Table 2). Four species were only found in
layers. the wild part of the pool: Juncus bufonius L.,
To quantify seedling dynamics, the cumulative Echinochloa crus-galli L., Ranunculus sardous C.,
number of seedlings found in each pot for each Elatine alsinastrum L. Seedlings abundance of a
species recorded among the 6 week sampling given species present in both parts of the pool
assessments was considered as a dependent var- differed (e.g. P. persicaria was more abundant in
iable. To identify distribution trends of number of the cultivated part while J. tenageia was more
seedlings and species richness across the upper abundant in the wild part).
(U) and the lower (L) sediment layer within each All together, more seedlings emerged from the
pool (Ec, Ew and F), we performed t-test for lower layers of the fallow pools (t = –5.46,
paired comparisons. For these analyses, the d.f. = 14, P < 0.001 for Ew and t = –3.13,
numbers of seedlings and species richness were d.f. = 14, P = 0.007 for F). But no difference
log transformed. We used S-PLUS (Math Soft was found between layers in the cultivated part
1999) for all our statistical analysis and consid- (t = –0.08, d.f. = 14, P = 0.93) of the pool
ered a test as statistically significant at 5% level. (Fig. 2a). With regards to the number of different

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Fig. 1 Relationship (a) Not Cultivated-F (b) Not Cultivated-Ew (c) Cultivated-Ec
between number of seeds 8
of the whole community
(log transformed) and 6
depth in each pool. Seeds

Seeds
were directly counted 4
after the sieving of each
depth layer from (a) the 2
unploughed F pool, (b)
the uncultivated area
0
Ew and (c) the cultivated
0 3 6 9 12 15 0 3 6 9 12 15 0 3 6 9 12 15
area Ec
Depth (cm)

Table 2 Total number of seedlings of each wetland taxum d.f. = 14, P = 0.0115), but no difference between
counted in all pots during the direct seedling experiment layers in fallow pools was detected (t = –1.97,
Species Ec Ew F d.f. = 14, P = 0.068 for Ew and t = –0.12,
d.f. = 14, P = 0.90 for F) (Fig. 2b).
Lythrum hyssopifolia 347 576 534
Lythrum portula 2733 2735 20
Juncus tenageia 62 1045 62
Focus on Damasonium alisma
Polygonum persicaria 170 15 36
Damasonium alisma 527 598 355 The number of D. alisma seeds per soil sample
Gnaphalium uliginosum 2 19 74 from sieved seed bank differed among pools
Juncus bufonius 0 60 34
Echinochloa crus-galli 0 6 1380
(P = 0.005) (Table 3). In particular, the number
Ranunculus sardous 0 8 36 of seeds showed an exponential decrease with
Elatine alsinastrum 0 3 0 depth in the uncultivated pools whereas there was
a slight linear increase of seed quantity with depth
in the ploughed pool (Fig. 3). Germination per-
species of seeds that germinated in the pots, more centage was also different among pools according
species in the lower layer compared to the upper to the depth at which they were extracted. In the
one were found in the cultivated pool (t = 2.90, cultivated pool, seeds from the deeper layers had

(a) Number of seedlings (b) Species richness

80 6

5
60
4
* * *
40 3

2
20
1

0 0
U L U L U L U L U L U L
Ec Ew F Ec Ew F

Fig. 2 (a) Seedlings distribution among layers revealed by standard deviation. An asterisk (*) denotes significant
direct seedling emergence experiment. Each sample was differences (P < 0.05) between U and L (t-test for paired
divided into two layers: U (the upper layer) in grey bars data S-PLUS 2000). (b) Relative distribution of species
and L (the lower layer) in black bars. Each bar represents richness recorded during seedling emergence for U and L
the mean cumulative data (log transformed) with its in each pool

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Table 3 Comparison of D. alisma seed numbers between 100

Germination percentage (%)

pools including the effect of pool (Ew, Ec and F), depth Ec ( ): r² =0.98; p<0.001
(five layers) and interaction between pool and depth 80

Source DF F Prob > F 60 Ew ( ): r² =0.92; p=0.03

Pool 2 5.50 0.005 40

Depth 1 70.53 <0.001
Pool · depth 2 32.89 <0.001 20 F ( ): r² =0.89; p=0.049
Core was considered as a random effect in the mixed 0
model (LME S-PLUS). Number of seeds was log 0 3 6 9 12 15
transformed for this analysis Depth (cm)

a higher germination percentage than those of the Fig. 4 Germination percentage of D. alisma seeds as a
function of the depth layer in each pool. Each value
upper layers. Conversely, in the wild pools, the represents, for a given depth, the percentage of seeds
seeds from the upper layers had a higher germi- (among 150) that germinated in the incubator room.
nation rate than the seeds from the deeper layers Square, triangle and ring symbol represent respectively
(Fig. 4). results for Ew, F and Ec

Since one of the aims of ploughing is to turn over

Discussion the soil, the quantitative distribution of soil seed
banks was deeply affected and seeds were mostly
Impact of ploughing on plant community buried by ploughing in cultivated pools.
We also found that the relative abundance of
Seed banks distribution was strongly affected by species among pools in the seedling emergence
ploughing and differed between cultivated and was contrasted: some species were more abun-
wild pools. In the wild pools, more seeds were dant in the cultivated part of the E pool while
counted from the samples cored at the surface of other species were mainly present in the wild part.
the soil than from deeper layers. An opposite For any given seed bank, the resulting emerging
trend was found in the cultivated pool. Concern- flora can be very different between pools because
ing the number of seeds that germinated from the for each species, a particular set of conditions
soil samples (seedling emergence), no germina- needs to be fulfilled in order to ensure successful
tion occurred in the sterilized soil pots. Therefore, germination and growth. In particular, in tempo-
we could assume that all the seedlings found in rary pools, the fluctuating level of water was
the pots during the experiment were the result of pointed out as a major factor influencing seedling
seeds germinating from samples cored in the recruitment from the seed bank (van der Valk
pools. Again, more seedlings emerged from the 1978). This seed bank property was seen as a
upper layers of the wild pools whereas no temporal storage effect because variable condi-
difference was observed among layers in the tions will lead to the recruitment of some species
cultivated pool. These first results were expected. only, excluding the others (Bonis 1998; Bliss and

Fig. 3 Relationship (a) Not Cultivated-F (b) Not Cultivated-Ew (c) Cultivated-Ec
between number of seeds 8
of D. alisma (log
transformed) and depth in 6
each pool. Seeds were
Seeds

counted directly after the 4

sieving of each depth
layer from (a) the 2
unploughed F pool, (b)
the uncultivated area Ew
0
and (c) the cultivated area 0 3 6 9 12 15 0 3 6 9 12 15 0 3 6 9 12 15
Ec
Depth (cm)

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Zedler 1998; Nicol et al. 2003; Warwick and Bonis and Lepart 1994; De Winton et al. 2000).
Brock 2003). This property induces a contrast But more interestingly, for the pool located in
between species’ abundance in the seed banks the cultivated field, we showed that the opposite
and species abundance observed in the field tendency was produced: seeds were more abun-
among adult plants (Jutila 2003). Other agricul- dant and had a better germination rate in the
tural practices such as the use of herbicide or deeper soil layers. This is probably partly due to
fertilisers may contribute to shape the seedlings’ the interaction between the life cycle of the
recruitment as well, and therefore certainly con- species and the timing of ploughing. Plants
tribute to the selection of a different community release seeds in late spring and the soil is
from the seed bank (Tørresen and Skuterud 2002; ploughed during fall, burying the seeds that
Shaukat and Siddiqui 2004). have been produced a few months before, and
Interestingly, we found not only that more bringing up old seeds that had been produced
species were located deep in the soil in the formerly (one or more years before). Since soil
cultivated pool, but also that most of their seeds samples were cored during early springtime, the
were actually still viable. This highlights that best seeds (the youngest ones) were located
many species of these wetland communities are deep in the soil. Therefore, ploughing had not
able to bear a certain period of burying. In fact, only accumulated most of the seeds in deep
a relatively large proportion of rare and endan- layers, but had also concentrated the best seeds.
gered species living in temporary pools are Beyond this temporal aspect, which likely
expected to form long-term persistent seed explains the inversion of the seed quality pattern
banks, because this trait is susceptible to be in ploughed pool, the moist conditions probably
selected in unpredictable environmental changes also enhance seed survival in deep layers.
(Hölzel et al. 2004). Consequently, for this kind Indeed, for this semi-aquatic species, dormant
of ephemeral wetland community, most seeds seeds remain dormant as long as they are either
are not killed by ploughing. Instead, a large part constantly damp or constantly submerged (Bir-
of the seed bank is stored in deep soil layers. As kenshaw 1994). These first results concerning D.
these results suggest, in storing seeds in the deep alisma are consistent with the spatial storage
layers of the soil, ploughing induces a spatial effect we have mentioned for the community
storage effect on the seed banks of such results.
communities. Birkenshaw (1994) showed that for D. alisma,
seeds can germinate only when situated in the
Impact of ploughing on Damasonium alisma first 3 cm of the soil. Consequently, seeds that
were actually able to germinate in our study were
We also focussed on a rare threatened species, those of the first layer (i.e the first 3 cm). As our
D. alisma, that is typically found in temporary results suggested, these seeds were more abun-
wetlands. As for community results, the vertical dant and had a better germination rate in the wild
distribution of seeds was strongly affected by pools. Therefore, one should expect to find more
ploughing; for the wild pools, the seed distribu- adult plants in the wild pools, than in ploughed
tion showed an exponential decrease with depth, ones. Yet the botanical inventory performed in
whereas a slight linear increase of seed abun- 2002 showed that D. alisma adult plants were
dance with depth was observed in the cultivated about three times more abundant in the culti-
pool. Moreover, our results concerning germina- vated part than in the wild part of the pool
tion percentage showed that, in the wild pools, (Decencière 2002). Moreover, this species is
the seeds located in the upper layers (the first known for being chiefly present in disturbed
3 cm) had a higher germination rate than the pools (Arnal 1996). At first sight, these field
seeds in the deeper layers (from 3 to 15 cm). results could be surprising according to our seed
This result is consistent with other seed bank bank investigation, since we expected to find
studies of other species performed in temporary more adult plants in the wild pools. Thus, the
pools, though uncultivated (Grillas et al. 1993; quantity and quality of the seeds are not the only

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52 Plant Ecol (2007) 192:45–53

factors that influence the relative abundance of References

the species among pools. It is likely that in wild
pools, post germination obstacles such as compe- Arnal G (1996) Les plantes protégées d’Ile-de-France.
Parthénope Collection. Biotope, Paris
tition with other species affect seedlings growth. Adams VM, Marsh DM, Knox JS (2005) Importance of
In cultivated pools, perennial herbs were re- the seed bank for population viability and population
moved by ploughing, competition was probably monitoring in a threatened wetland herb. Biol Con-
lower and thus the viability of D. alisma seedlings serv 124:425–436
Beuret E (1989) Influence des pratiques culturales sur
was enhanced. l’évolution de la flore adventice: étude du potentiel
D. alisma seeds seemed to tolerate the bury- semencier des sols. Revue suisse agricole 21:75–82
ing conditions since the seeds deeply cored had Birkenshaw CR 1994 Aspects of the ecology and conser-
a higher germination ability. When buried, seeds vation of Damasonium alisma Miller in Western
Europe. Watsonia 20:33–39
are protected from unfavourable environmental Bliss SA, Zedler PH (1998) The germination process in
conditions while in the wild pools, most of the vernal pools: sensitivity to environmental conditions
seeds germinate at the same time. Such conclu- and effects on community structure. Oecologia
sions agree with demographic models showing 113:67–73
Bonis A (1998) Coexistence in a variable and unpredict-
that seed survival is crucial for population able habitat: the importance of the seed bank and
viability of wetland plants (Adams 2005). Dur- contrasted regeneration niche for annual plants in
ing a drought, for example, most of the seedlings temporary pools. Arch Hydrobiol Spec Issues Advanc
would be killed in wild pools, whereas seeds are Limnol 52:53–69
Bonis A, Lepart J (1994) Vertical structure of seed bank
largely protected in deep layers in cultivated and the impact of depth of burial on recruitment in
fields and are able to give birth to new seedlings two temporary marches. Plant Ecol 112:127–139
the following season, after the next ploughing. Cohen D (1966) Optimizing reproduction in a random
In addition, seedlings will face lower competi- varying environment. J Theoret Biol 12:119–129
Danton P, Baffray M (1995) In: Rocher Y (ed) Inventaire
tion. In that manner, as for the whole commu- des plantes protégées en France. AFCEV, Nathan
nity, seed banks constitute a spatial protected Decencière B (2002) Les mouillères de l’Essonne: Ap-
stock of individuals at the population level for proche écologique et ethnologique. Rapport de stage
D. alisma. de DEA. Conservatoire botanique national du Bassin
Parisien. Master report. MNHN, Paris, France
In conclusion, we showed that ploughing had a De Winton MD, Clayton JS, Champion PD (2000)
strong effect on species distribution and abun- Seedling emergence from seed banks of 15 New
dance of temporary wetland communities. More- Zealand lakes with contrasting vegetation histories.
over, in allowing constitution and preservation of Aquat Bot 66:181–194
Gallego-Fernández JB, Garcia-Mora MR, Garcia-Novo F
seed banks and in removing perennial species, (1999) Small wetland lost: biological conservation
rare annual species such as D. alisma can be hazard in mediterranean landscapes. Envir Conserv
protected in the long run. In this particular case, 26:190–199
we believe that agricultural practices could have a Ghersa CM, Martinez-Ghersa MA (2000) Ecological
correlates of weed seed size and persistence in the
beneficial effect for the preservation of rare soil under different tilling systems: implications for
species. Further investigations are needed, how- weed management. Field crops Res 67:141–148
ever, to understand the effects of other agricul- Grillas P, Garcia-Murillo P, Geertz-Hansen O, Marbá N,
tural practices (Hölzel et al. 2004). The Montes C, Duarte CM, Tan Ham L, Grossmann A
(1993) Submerged macrophyte seed bank in a Med-
understanding of the processes that maintain this iterranean temporary marsh: abundance and relation-
specific flora in the pools should be taken into ship with established vegetation. Oecologia 94:1–6
account in order to develop more sustainable Hedrick PW (1995) Genetic polymorphism in a temporally
management policies. varying environment: effects of delayed germination
or diapause. Heredity 75:164–170
Hölzel N, Otte A (2004) Assessing soil seed bank
Acknowledgements We would like to thank Gerard
persistence in flood-meadows: the search for reliable
Arnal for floristic inventories, Beatriz Decencière for
traits. J Veg Sci 15:93–100
providing data and assistance in the field and Dominique
Jalili A, Hamzeh’ee B, Asri Y, Shirvany A, Yazdani S,
Debray and Gregoire Devictor for manuscript
Khoshnevis M, Zarrinkamar F, Ghahramani M,
improvement.
Safavi R, Shaw S, Hodgson JG, Thompson K,

123
Plant Ecol (2007) 192:45–53 53

Akbarzadeh M, Pakparvar M (2003) Soil seed banks Standifer LC (1980) A technique for estimating weed seed
in the Arasbaran protected area and their signifi- populations in cultivated soil. Weed Sci 28:134–138
cance for conservation management. Biol Conserv Stoate C, Boatman ND, Borralho RJ, Carvalho CR, De
109:425–431 Snoo GR, Eden P (2001) Ecological impacts of arable
Jutila HM (2003) Germination in Baltic coastal wetland intensification in Europe. J Env Manage 63:337–365
meadows: similarities and differences between vege- Stöcklin J, Fisher M (1999) Plants with longer-lived seeds
tation and seed bank. Plant Ecol. 166:275–293 have lower local extinction rates in grassland rem-
Kalisz S (1991) Experimental determination of seed bank nants 1950–1985. Oecologia 120:539–543
age structure in the winter annual Collinsia verna. Templeton AR, Levin DA (1979) Evolutionary conse-
Ecology 72:575–585 quences of seed pools. Ame Natura 114:232–249
Lefeuvre J-C, Fustec E, Barnaud G (2000) De l’élimina- Thrall PH, Pacala SW, Silander JA (1989) Oscillatory
tion à la reconquête des zones humides. In: Fustec E, dynamics in populations of an annual weed species
Lefeuvre J-C et al (eds) Fonction et valeurs des zones Abutilon theophrasti. J of Ecol 77:1135–1149
humides. Dunod, Paris, 426 pp Tørresen KS, Skuterud R (2002) Plant protection in spring
Luzuriaga AL, Escuderao A, Olano JM, Loidi J (2005) cereal production with reduced tillage. IV. Changes in
Regenerative role of seed banks following an inten- the weed flora and weed seedbank. Crop Prot 21:179–
sive soil disturbance. Acta Oecologica 27:57–66 193
Math Soft 1999. S-PLUS 2000. Data analysis products van der Valk AG (1978) The role of seed banks in the
division, MathSoft, Seatlle. WA vegetation dynamics of prairie glacial marshes. Ecol-
Middleton BA (2003) Soil seed banks and the potential ogy 59: 322–335
restoration of forested wetlands after farming. J Appl Vitalis R, Glemin S, Olivieri I (2004) When genes go to
Ecol 40:1025–1034 sleep: the population genetic consequences of seed
Miller GR, Cummins RP (2003) Soil seed banks of dormancy and monocarpic perenniality. Ame Natura
woodland, heathland, grassland, mire and montane 163:295–311
communities, Cairngorm Mountains, Scotlan. Plant Vuille FL (1987) Reproductive biology of the genus
Ecol 168:255–266 Damasonium (Alismataceae). Pl Syst Evol 157:63–71
Nicol JM, Ganf GG, Pelton GA (2003) Seed banks of a Warr SJ, Thompson K, Kent M (1993) Seed banks as a
southern Australian wetland: the influence of water neglected area of biogeographic research: a review of
regime on the final floristic composition. Plant Ecol literature and sampling techniques. Prog Phys Geog
168:191–205 17:329–347
Pacala E (1986) Neighbourhood models of plant popula- Warwick NWM, Brock MA (2003) Plant reproduction in
tion dynamics. 2. Multi-species models of annuals. temporary wetlands: the effects of seasonal timing,
Theoret Popul Biol 29:262–292 depth, and duration of flooding. Aquat Bot 77:153–
Pinhero JC, Douglas MB (2000) Mixed-effects models in S 167
and S-PLUS. Springer, New-York Wheeler BR (2000) Starfruit Damasonium alisma. Pro-
Roberts HA (1981) Seed banks in soils. Adv Appl Biol gress report on the (1999) project. Report No. 146.
6:1–55 London: Plantlife
Shaukat SS, Siddiqui IA (2004) Spatial pattern analysis of Wienhold CE, van der Valk AG (1989) The impact of
seeds of an arable soil seed bank and its relationship duration of drainage on the seed banks of northern
with above-ground vegetation in an arid region. prairie wetlands. Can J Bot 67:1878–1884
J Arid Env 57:311–327

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