1. No category

Ekológia (Bratislava)
Vol. 24, Supplement 1/2005, p. 7–27
DIASPORES – POTENTIAL OR REAL POWER OF
WILD PLANTS? LIFE CYCLE
BOŽENA ŠERÁ
Institute of Systems Biology and Ecology, Academy of Sciences of the Czech Republic, Na Sádkách 7,
370 05 České Budějovice, The Czech Republic, e-mail: [email protected]
Abstract
Šerá B.: Diaspores – potential or real power of wild plants? Life cycle. Ekológia (Bratislava),
Vol. 24, Supplement 1/2005, p. 7–27.
All organisms have only got a limited amount of energy in their life history. An investment into
reproduction is adequate to the growth condition of individual or population. For the plant survival
it is necessary to maximize the chances of the offspring to survive in all phases of development
(e.g. phase of seed). If seed/diaspore weight is relatively constant (interspecies variability), then
it is number of seeds, which is an important factor for growth and survival of population in
natural conditions (dispersal in the space and in the time). Number of seeds seems to be the
crucial traits for reproductive strategy of plants.
Key words: life history, seed production, seed weight, trade-off
Introduction
The amount of produced seeds, their weight and size change both within and among species depending on many traits, the abiotic and biotic factors in the locality are the most
important ones (Francland, 1975; Thompson, 1984; Wardlaw, Dunstone, 1984). Both of
these traits indicating the reproductive strategy trend of species depend on the plant fitness,
its height (Hendrix, 1984), specific leaf area (Westoby et al., 1996), pollination success
(Colosi, Cavers, 1984; Boyd et al., 1990), time of pollination (Galen, Stanton, 1991), season and plant age (Downs, McQuilkin, 1944 sec. Begon et al., 1990; Cavers, Steele, 1984),
fruit position on the plant body (Waller, 1982; Stanton, 1984; Wulff, 1986), and seed position in fruit (e.g. Fabaceae; Salisbury, 1942). Both seed weight and number of seeds are
given genetically, but there is an enormous intraspecific and interspecific variability. Both
of these variabilities reflect common life history trade-off of species (Grime, 1977; Begon
et al., 1990; Šerá, Šerý, 2004).
Life history is a basic trait, which is in strong correlation to reproduction and to the
possibilities of the population growth. Plants are divided according to the number of flow7
ering per life into monocarps and polycarps. Monocarpic plants flower and produce seeds
once per their life, and polycarpic plants (perennials) produce seeds repeatedly. Monocarpic
plants are divided according to the length of their life into annuals, biennials (they flower in
the second year of their life), and monocarpic perennials yielding in the last year of their
life. There is a common principle, that there is a decrease in number of produced seeds
depending on the plant life span (Harper, 1977).
There is still one question about the general reproductive strategy of an organism (plant),
because the number of offspring (seeds) is only a part of the problem. The crucial importance for all organisms (plants) is what size offspring (seeds/diaspores) is produced. In
addition, many plants compared to animals are able to reproduce themselves not only by
a sexual propagation but in a vegetative way, too. What is the role of size and number of
seeds for plants, which are able of vegetative propagation? All the mentioned factors (life
span, number of flowering, lateral spread, seed/diaspore weight, number of produced seeds/
diaspores) participate in reproductive strategy of each of plant species in different measure.
Materials and methods
The number of diaspores produced by herbaceous vegetation (reproductive capacity of population) and diaspore
weight (mean of seed weight) were taken from Šerá and Šerý (2004). The data of herbaceous species from
Central Europe were complemented with life cycle traits and tendencies to lateral spread (Hejný, Slavík, 1988,
1990, 1992; Rothmaler et al., 1987; Slavík, 1995, 1997; Frank, Klotz, 1988; Kubát et al., 2002).
1. Number of flowering – monocarps, polycarps
2. Life span – annuals, biennials (and monocarpic perennials), perennials
3. Lateral spread – plants without lateral spread, with the ability of vegetative propagation to 0.30 m per year,
with the ability of vegetative propagation over 0.31 m per year.
This work understands „diaspore“ as a functional reproductive part (e.g. achene, grain, seed with a caruncula,
achene with a flying apparatus) (Cornelissen et al., 2003; Šerá, Šerý, 2004).
The data were calculated using the statistic programme STATISTICA (’99). The quantitative variables (reproductive capacity of population, mean of diaspore weight) were log transformed prior to the analysis to achieve their
normal distribution. All the statistic procedures were subsequently calculated from this transformed data set.
The differences among individual types of traits with regard to the number and weight of seeds were calculated using a multiple range analysis by the Tukey HSD test (for unequal n) at the significant level α = 0.05
(Table 1). Box-whisker plots were used for visualisation of these significant results (Fig. 1). Biennials and
monocarpic perennials (4 species) were calculated altogether, they are labelled as biennials (Table 1, Fig. 1).
Spearman correlation among qualitative (number of flowering, life span, lateral spread) and quantitative
traits (see above) was used (R = Spearman’s correlation coefficient), but Pearson correlation was counted only
between quantitative traits (r(X,Y) = Pearson’s correlation coefficient) (Table 2).
Results
Monocarpic species produced significantly more diaspores per year than polycarps. Similarly, annual species produced more diaspores than biennials (altogether with monocarpic
perennials) and biennials produced more diaspores than perennials. Plants without the veg8
T a b l e 1. Number of diaspores (reproductive capacity of population) and diaspore weight comparisions for
life history traits of herbaceous species. Traits followed by the same letters were not significantly different in
ANOVA analysis (Tukey HSD test for unequal n, log transformation, P < 0.05).
Traits
Reproductive capacity of Number of
flowering
population (1 000/m )
2
Life span
n
Diaspore weight (mg)
Life span
Media
n
SD
Monocarpsa
175
540
116
1 469
Polycarpsb
319
500
59
1 861
Annualsa
Biennials and Monocarpic
perennialsab
131
595
120
1 637
44
377
99
772
Perennialsb
319
500
59
1 861
Lateral spread Withouta
Number of
flowering
Mean
224
659
107
1 930
< 0.30 m per yearb
176
384
68
1 225
t 0.31 m per yearb
92
408
76
2 010
Monocarpsa
169
2.03
0.68
4.42
Polycarpsa
286
2.88
0.65
8.98
Annualsa
Biennials and Monocarpic
perennialsa
126
1.79
0.58
4.13
43
2.72
1.07
5.17
Perennialsa
286
2.88
0.65
8.98
Lateral spread Withouta
< 0.30 m per yeara
219
3.10
0.68
9.80
157
2.21
0.70
4.95
t 0.31 m per yeara
79
1.79
0.58
4.36
etative propagation ability produced more diaspores than species with the lateral spread.
On contrary to all expectation, no significant difference was found in the traits with regard
to the mean weight of diaspores. The calculated results are summarised and visualised in
the Table 1 and Fig. 1.
Many correlations among quantitative and qualitative traits were logical (positive correlation between number of flowering and lateral spread, and life span; positive correlation
between life span and lateral spread) (Table 2). Some relations were interesting for explanation of diaspore production in relation to plant reproductive strategy (see Discussion).
Discussion
The results of this work confirmed, that the number of produced diaspores decreased with
the plant life span, with the number of reproduction possibilities per life, and with the
ability of lateral spread (Fig. 1; Šerá, Šerý, 2004). A huge production of offspring is strate9
7,0
8
6,5
Reproductive capacity of population
Reproductive capacity of population
7
6,0
5,5
5,0
4,5
4,0
3,5
6
5
4
3
3,0
2,5
2
Monocarps
Polycarps
Number of flowering
Annuals
Biennials
Perennials
Life span
8
Reproductive capacity of population
7
6
5
4
3
2
0
1
Lateral spread
2
Min-Max
25%-75%
Median value
Fig. 1. Box-whisker plots on significant differences
Tukey HSD test (for unequal n) among reproductive
strategy traits. Plots represent median, interval between
lower and upper quartiles, minimum and maximum
values of tested traits types. Details see in text.
gic for a monocarpic plant, reproducing itself only once per year than for a plant with
repeated flowering, which must invest into the persistent vegetative body parts (Salisbury,
1942). Annual plants are typical for temporary localities and a huge number of diaspores
helps them to survive successfully on the site (seed dispersal) and in time (seed bank)
(Southwood, 1988; Grime, 2001).
On contrary, a polycarpic plant flowering more times per its life must invest not only to
the reproduction but to the vegetative growth, too. The allocation to repeated flowering is
balanced with the probability of the survival of the parent plant (e.g. Šerá, 1995). Polycarpic
plant (perennials) must be more competitive to other species growing on the same site than
plants with the shorter life cycle.
Annual and polycarpic perennial plants have got lighter diaspores in average than
polycarpic plants (Baker, 1972; Silvertown, 1981) and biennials have got heavier diaspores
than annual and perennial plants (Lovett Doust, 1980; Thompson, 1984). No such claims
were confirmed, because no significant difference was proved among the investigated groups
in relation to the diaspore weight (Tables 1, 2). A number of diaspores appears to be a more
important trait for the reproductive strategy of plants than a diaspore weight. A diaspore
weight is less various than their number (Harper, 1977; Šerá, Šerý, 2004). Experiments
10
Number of flowering
1.000
Life span
****0.982
1.000
Lateral spread
****0.607
****0.630
1.000
-0.006
0.007
-0.024
1.000
*** - 0.197
**** - 0.200
**- 0.171
**- 0.760
Diaspore weight
Reproductive capacity of
population
Reproductive capacity of population
Diaspore weight
Lateral spread
Life span
Number of flowering
T a b l e 2. Matrix of trates correlation on life history plants. Spearman correlation was used between
qualitative trates and Pearson correlation between quantitative. R and r(X,Y) were counted, P < 0.01*,
P < 0.001**, P < 0.0001***, P < 0.00001****. See the text.
1.000
running in various growing conditions confirmed, that diaspore weight is phenotypically
very constant for many plant species (e.g. Fenner, 1985). The number of produced diaspores
is more variable comparing to the diaspore weight (e.g. Šerá, Šerý, 2004). The most important traits influencing on total number of produced diaspores are biotical and abiotical
conditions on the locality. In bad natural conditions, a fruit-bearing plant sacrifices some
of the diaspores rather than their weight or size (Lloyd, 1987).
Flowering plants may reproduce in two ways: through seeds/diaspores (largely sexually)
or by means of vegetative organs (asexually – offshoots, tillers, bulbs, creeping rhizomes,
etc.). Some plant species may reproduce only by diaspores (usually annuals), or entirely in
a vegetative way (e.g. aquatic plants). However, most of the plants use both ways, namely
° rgen, Willson, 1994).
in a dependence on specific life conditions (e.g. A
An investment to the diaspores is favourable for source/energy distribution in the plant
body, than allocation to vegetative reproduction, which is more energetically demanding.
Diaspores are considerably smaller than the parent plant consequently they may be produced in a huge number. In addition, small diaspores can be well dispersed (by wind, animals, water) to long distances from the parent plant (Pijl, 1982; Lhotská et al., 1987). Such
species may quickly expand to new sites. They are successful in young open communities
11
(Hodgson, Mackey, 1986; Grime, Hillier, 1992; Grime, 2001). Diaspores may survive bad
condition in the seed bank (e.g. drought) in contrast to vegetative offspring. Such thoughts
are supported by the fact that the number of produced diaspores was several higher in
species without lateral spread than in species with vegetative propagation (Tables 1, 2).
The benefits of vegetative propagation are above all in fast offspring growth, their smaller
mortality, possibility to locomotion, and in bigger ability of competition (Silvertown, Lovett
Doust, 1993). The species which is able of lateral spread is not vitally dependent on reproduction by diaspores and consequently the number of produced diaspores is lower, than the
production of species with reproduction only by diaspores (Tables 1, 2).
The reproduction strategy (seeds/diaspores) is one of the crucial points in final species
composition and vegetation succession (Jakrlová, 1985; Rosenzweig, 1995; Grime, 2001).
In addition, the plant reproduction contributes to historical change of plant communities or
habitats (e.g. alien and weed plants – Crawley, 1987; Pyšek et al., 2005). The number of
diaspore is an important factor of the population viability; consequently it may seriously
cause all plant communities.
Translated by author and D. Simek
Acknowledgement
I am obliged to D. Simek for her linguistic revision of the paper. A funding was provided by grant No. 1P05OC049
and by Institutional research plan No. AV0Z60870520.
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13
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Received 25. 7. 2005
Šerá B.: Diaspory – potenciální nebo reálná síla divokých rostlin? Životní cyklus.
Každý organismus má pro svůj životní cyklus k dispozici jen limitované množství energie. Energetické příděly
do reprodukce odpovídají růstové kondici jedince/populace. Pro přežívání jedinců/populace rostlin je nezbytné
maximalizovat šanci přežití semenáčků a to ve všech fázích přežívání (např. semena). Pokud je hmotnost semene
relativně konstantní (vnitrodruhová variabilita), potom je počet semen důležitým faktorem pro růst a přežívání
celé populace v přírodních podmínkách (šíření v prostoru a v čase). Počet semen se zdá být klíčovým faktorem
pro úspěšnou reprodukci rostlin.
14
15
Achillea millefolium L.
Achillea ptarmica L.
Aconitum variegatum L.
Actaea spicata L.
Aethusa cynapium L.
Agrimonia procera W a l l r.
Agrostis canina L.
Agrostis capillaris L.
Agrostis stolonifera L.
Ajuga genevensis L.
Ajuga reptans L.
Alchemilla monticola O p i z
Alisma plantago–aquatica L.
Alliaria petiolata (M. B i e b.) C a v a r a et G r a n d e
Allium ursinum L.
Alopecurus pratensis L.
Alyssum alyssoides (L.) L.
Amaranthus retroflexus L.
Anagallis arvensis L.
Anemone nemorosa L.
Angelica sylvestris L.
Anthemis arvensis L.
Anthoxanthum odoratum L.
Anthriscus sylvestris (L.) H o f f m .
Apera spica–venti (L.) P. B e a u v.
Aphanes arvensis L.
Arabidopsis thaliana (L.) H e y n h.
Arabis glabra (L.) B e r n h.
Arabis hirsuta (L.) S c o p.
Arctium lappa L.
Arctium tomentosum M i l l.
Arenaria serpyllifolia L.
Arnica montana L.
Arrhenatherum elatius (L.) J. P r e s l et C. P r e s l
Artemisia vulgaris L.
Name
Asteraceae
Asteraceae
Ranunculaceae
Ranunculaceae
Apiaceae
Rosaceae
Poaceae
Poaceae
Poaceae
Lamiaceae
Lamiaceae
Rosaceae
Alismataceae
Brassicaceae
Liliaceae
Poaceae
Brassicaceae
Amaranthaceae
Primulaceae
Ranunculaceae
Apiaceae
Asteraceae
Poaceae
Apiaceae
Poaceae
Rosaceae
Brassicaceae
Brassicaceae
Brassicaceae
Asteraceae
Asteraceae
Caryophyllaceae
Asteraceae
Poaceae
Asteraceae
Family
achene
achene
seed
seed
mericarp
achene
caryopsis
caryopsis
caryopsis
nutlet
nutlet
achene
achene
seed
seed
caryopsis
seed
seed
seed
achene
mericarp
achene
caryopsis
mericarp
caryopsis
achene
seed
seed
seed
achene
achene
seed
achene
caryopsis
achene
Diaspore
Reproductive
capacity
of population
101 882
66 400
35 122
1 164
126 314
9 265
385 148
491 234
62 463
124 933
53 753
55 413
330 444
138 508
4 751
90 309
103 913
138 123
212 971
60 182
15 643
114 100
211 895
62 448
250 801
242 205
4 448 014
1 483 500
1 103 333
15 167
29 117
1 256 927
67 304
25 151
177 548
Diaspore
weight
(mg)
0.033
–
1.907
5.895
1.600
2.000
0.047
0.056
0.045
–
0.730
0.221
0.250
2.090
3.365
0.520
0.381
0.409
0.256
1.283
2.500
0.130
0.286
2.033
0.141
0.172
0.012
0.014
0.074
13.378
11.130
0.025
0.955
2.130
0.090
per life
PC
PC
PC
PC
MC
PC
PC
PC
PC
PC
PC
PC
PC
MC
PC
PC
MC
MC
MC
PC
MC
MC
PC
PC
MC
MC
MC
MC
MC
MC
MC
MC
PC
PC
PC
of flowering
Number
P
P
P
P
A
P
P
P
P
P
P
P
P
A
P
P
A
A
A
P
B
A
P
P
A
A
A
A
B
B
B
A
P
P
P
span
Life
1
1
1
0
0
1
2
2
2
2
2
1
2
0
0
1
0
0
0
2
1
0
1
0
0
0
0
0
0
0
0
0
0
1
1
spread
Lateral
Appendix: Ground data about number of produced diaspores (reproductive capacity of population), diaspore weight, height of plant, number of flowering per
life (MC – monocarpic plants, PC – polycarpic plants), life span (A – annuals, B – biennials and monocarpic perennials, P – perennials), and lateral spread
(0 – without vegetative propagation, 1 – propagation to 0.30 m per year, 2 – propagation over 0.31 m per year). Details see in text.
16
Name
Aruncus vulgaris R a f i n.
Asarum europaeum L.
Astragalus glycyphyllos L.
Astrantia major L.
Atriplex patula L.
Atriplex sagittata B o r k h.
Atropa bella–dona L.
Avenella flexuosa (L.) P a r l.
Ballota nigra L.
Barbarea vulgaris R. B r.
Batrachium aquatile (L.) D u m o r t
Bellis perennis L.
Berteroa incana (L.) D C.
Betonica officinalis L.
Bidens cernua L.
Bidens frondosa L.
Bidens tripartita L.
Brachypodium pinnatum (L.) P. B e a u v.
Brachypodium sylvaticum (H u d s.) P. B e a u v.
Brasica napus L.
Briza media L.
Bromus erectus H u d s.
Bromus hordeaceus L.
Bromus inermis L e y s s
Bromus sterilis L.
Bromus tectorum L.
Bryonia alba L.
Butomus umbellatus L.
Calamagrostis arundinacea (L.) R o t h
Calamagrostis canescens (W e b e r) R o t h
Calamagrostis epigejos (L.) R o t h
Calamagrostis villosa (C h a i x ex V i l l.) J. F. G m e l
Callitriche palustris agg.
Calluna vulgaris (L.) H u l l
Caltha palustris L.
Calystegia sepium (L.) R. B r.
Camelina microcarpa A n d r z.
Campanula patula L.
Campanula persicifolia L.
Appendix. (Continued)
Rosaceae
Aristolochiacea
Fabaceae
Apiaceae
Chenopodiaceae
Chenopodiaceae
Solanaceae
Poaceae
Lamiaceae
Brassicaceae
Ranunculaceae
Asteraceae
Brassicaceae
Lamiaceae
Asteraceae
Asteraceae
Asteraceae
Poaceae
Poaceae
Brassicaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Cucurbitaceae
Butomaceae
Poaceae
Poaceae
Poaceae
Poaceae
Callitrichaceae
Ericaceae
Ranunculaceae
Convolvulaceae
Brassicaceae
Campanulaceae
Campanulaceae
Family
follicle
seed
seed
mericarp
achene
achene
seed
caryopsis
nutlet
seed
achene
achene
seed
nutlet
achene
achene
achene
caryopsis
caryopsis
seed
caryopsis
caryopsis
caryopsis
caryopsis
caryopsis
caryopsis
seed
seed
caryopsis
caryopsis
caryopsis
caryopsis
seed
seed
seed
seed
seed
seed
seed
Diaspore
Reproductive
capacity
of population
32 459
8 467
7 221
41 293
67 945
40 959
32 613
81 857
91 670
212 286
88 204
67 407
139 388
109 926
53 041
53 308
231 371
65 000
11 131
180 472
30 956
120 964
63 610
27 077
67 835
105 400
1 619
124 796
15 887
295 597
258 327
34 102
148 290
761 890
245 794
969
373 133
2 375 000
375 000
Diaspore
weight
(mg)
0.090
3.740
3.970
1.800
7.600
1.000
1.030
0.266
1.675
0.600
–
0.082
0.270
1.159
0.907
2.056
2.100
–
2.730
–
0.523
–
2.100
–
3.900
2.000
11.800
0.200
0.380
0.200
0.256
0.817
0.023
0.060
0.663
22.870
0.180
0.020
0.080
Number
per life
PC
PC
PC
PC
MC
MC
PC
PC
PC
MC
MC
PC
MC
PC
MC
MC
MC
PC
PC
MC
PC
PC
MC
PC
MC
MC
PC
PC
PC
PC
PC
PC
MC
PC
PC
PC
PC
MC
PC
of flowering
Life
P
P
P
P
A
A
P
P
P
B
A
P
A
P
A
A
A
P
P
A
P
P
A
P
A
A
P
P
P
P
P
P
A
P
P
P
P
B
P
span
2
1
1
0
0
0
0
1
0
0
2
0
0
0
0
0
0
2
1
0
1
1
0
1
0
0
0
1
2
2
2
2
0
1
0
0
0
0
0
spread
Lateral
17
Name
Campanula rapunculoides L.
Campanula rotundifolia L.
Campanula trachelium L.
Capsella bursa–pastoris (L.) M e d i k.
Cardamine pratensis L.
Cardaminopsis halleri (L.) H a y e k
Carduus crispus L.
Carduus nutans L.
Carduus personata (L.) J a c q.
Carex brizoides L.
Carex canescens L.
Carex elata A l l.
Carex flacca S c h r e b.
Carex gracilis C u r t i s.
Carex hirta L.
Carex humilis L e y s s.
Carex nigra (L.) R e i c h a r d
Carex ovalis G o o d e n.
Carex pairaei F. W. S c h u l t z
Carex pallescens L.
Carex panicea L.
Carex pilosa S c o p.
Carex praecox S c h r e b.
Carex riparia C u r t i s
Carex rostrata S t o k e s ex W i t h.
Carex spicata H u d s.
Carex vesicaria L.
Carex vulpina L.
Carlina acaulis L.
Carum carvi L.
Centaurium erythraea R a f n
Cerastium arvense L.
Cerastium holosteoides F r.
Cerastium pumilum C u r t.
Chaerophyllum aromaticum L.
Chaerophyllum aureum L.
Chaerophyllum hirsutum L.
Chaerophyllum temulum L.
Chamerion angustifolium (L.) H o l u b
Appendix. (Continued)
Campanulaceae
Campanulaceae
Campanulaceae
Brassicaceae
Brassicaceae
Brassicaceae
Asteraceae
Asteraceae
Asteraceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Cyperaceae
Asteraceae
Apiaceae
Gentianaceae
Caryophyllaceae
Caryophyllaceae
Caryophyllaceae
Apiaceae
Apiaceae
Apiaceae
Apiaceae
Onagraceae
Family
seed
seed
seed
seed
seed
seed
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
mericarp
seed
seed
seed
seed
mericarp
mericarp
mericarp
mericarp
seed
Diaspore
Reproductive
capacity
of population
175 000
2 143 146
268 837
1 965 033
236 918
3 521 393
21 200
3 498
6 113
44 620
55 143
22 615
12 080
27 863
38 232
14 000
118 629
48 912
40 776
64 256
37 269
9 761
52 274
23 261
26 575
27 900
29 776
17 967
29 556
95 191
12 252 500
560 841
545 067
3 061 602
79 401
77 067
15 184
86 914
2 930 167
Diaspore
weight
(mg)
0.220
0.030
0.143
0.113
0.162
–
1.902
1.968
2.540
0.260
0.153
0.835
0.886
0.450
0.910
1.780
1.643
0.274
0.080
1.230
0.985
1.000
–
–
0.563
1.700
0.543
1.430
3.460
1.050
0.004
0.116
0.091
0.035
11.200
7.300
2.100
10.400
0.104
Number
per life
PC
PC
PC
MC
PC
PC
MC
MC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
MC
MC
PC
PC
MC
PC
PC
PC
MC
PC
of flowering
Life
P
P
P
A
P
P
B
B
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
B
A
P
P
A
P
P
P
B
P
span
1
1
2
0
0
1
1
1
1
2
2
1
2
2
2
1
2
1
1
1
2
1
1
2
2
1
2
1
0
0
0
1
0
0
1
0
1
0
2
spread
Lateral
18
Name
Chelidonium majus L.
Chenopodium album L.
Chenopodium bonus–henricus L.
Chenopodium glaucum L.
Chenopodium hybridum L.
Chenopodium pedunculare B e r t o l.
Chenopodium polyspermum L.
Chenopodium rubrum L.
Chenopodium strictum R o t h
Chenopodium suecicum M u r r
Cichorium intybus L.
Cicuta virosa L.
Circaea luteciana L.
Cirsium arvense (L.) S c o p.
Cirsium heterophyllum (L.) H i l l
Cirsium oleraceum (L.) S c o p.
Cirsium palustre (L.) S c o p.
Cirsium vulgare (S a v i) T e n.
Clinopodium vulgare L.
Colchicum autumnale L.
Conium maculatum L.
Consolida regalis G r a y
Convallaria majalis L.
Convolvulus arvensis L.
Conyza canadensis (L.) C r o n g u i s t
Coronilla varia L.
Corydalis cava (L.) S c h w e i g g. et K ö r t e
Corydalis solida (L.) C l a i r v.
Crepis biennis L.
Cruciata glabra (L.) E h r e n d.
Cuscuta europaea L.
Cyanus segetum H i l l
Cymbalaria muralis P. G a e r t n., B. M e y. et S c h e r b.
Cynodon dactylon (L.) P e r s.
Cynoglossum officinale L.
Cynosurus cristatus L.
Dactylis glomerata L.
Dactylorhiza fuchsii (D r u c e) S o ó
Dactylorhiza majalis (R c h b.) P. F. H u n t et S u m m e r h
Appendix. (Continued)
Papaveraceae
Chenopodiaceae
Chenopodiaceae
Chenopodiaceae
Chenopodiaceae
Chenopodiaceae
Chenopodiaceae
Chenopodiaceae
Chenopodiaceae
Chenopodiaceae
Asteraceae
Apiaceae
Onagraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Lamiaceae
Liliaceae
Apiaceae
Ranunculaceae
Liliaceae
Convolvulaceae
Asteraceae
Fabaceae
Fumariaceae
Fumariaceae
Asteraceae
Rubiaceae
Cuscutaceae
Asteraceae
Scrophulariaceae
Poaceae
Boraginaceae
Poaceae
Poaceae
Orchidaceae
Orchidaceae
Family
seed
achene
achene
achene
achene
achene
achene
achene
achene
achene
achene
mericarp
seed
achene
achene
achene
achene
achene
nutlet
seed
mericarp
seed
seed
seed
achene
seed
seed
seed
achene
mericarp
seed
achene
seed
caryopsis
nutlet
caryopsis
caryopsis
seed
seed
Diaspore
Reproductive
capacity
of population
286 799
312 589
23 442
110 556
228 120
206 018
264 321
222 192
65 468
79 928
76 096
30 500
5 806
32 776
5 750
13 985
59 469
53 070
216 818
86 767
92 388
197 647
2 469
909
800 476
129 900
17 664
13 183
74 896
243 185
190 748
25 166
59 560
55 550
14 453
96 897
138 622
4 074 563
4 720 600
Diaspore
weight
(mg)
0.697
0.656
1.990
0.102
1.296
0.698
0.239
0.114
0.330
0.620
0.738
2.200
0.550
1.485
2.330
3.640
1.202
4.627
0.452
7.900
1.000
0.830
26.160
11.370
0.048
0.810
1.225
5.250
0.982
–
0.600
2.870
0.128
–
28.700
2.800
0.963
0.001
0.002
Number
per life
PC
MC
PC
MC
MC
MC
MC
MC
MC
MC
PC
PC
PC
PC
PC
PC
MC
MC
PC
PC
MC
MC
PC
PC
MC
PC
PC
PC
MC
PC
MC
MC
PC
PC
MC
PC
PC
PC
PC
of flowering
Life
P
A
P
A
A
A
A
A
A
A
P
P
P
P
P
P
B
B
P
P
B
A
P
P
A
P
P
P
B
P
A
A
P
P
B
P
P
P
P
span
1
0
1
0
0
0
0
0
0
0
0
1
2
2
2
2
1
2
1
0
0
0
2
2
0
2
0
0
2
1
0
0
2
2
0
1
1
0
0
spread
Lateral
19
Name
Danthonia decumbens (L.) D C.
Daucus carota L.
Deschampsia cespitosa (L.) P. B e a u v.
Descurainia sophia (L.) W e b b ex P r a n t l
Dianthus deltoides L.
Digitalis grandiflora M i l l.
Digitalis purpurea L.
Digitaria sanguinalis (L.) S c o p.
Dipsacus fullonum L.
Echinochloa crus–galli (L.) P. B e a u v.
Echium vulgare L.
Eleocharis palustris (L.) R o e m. et S c h u l t.
Elytrigia repens (L.) D e s v.
Epilobium ciliatum R a f.
Epilobium hirsutum L.
Epilobium montanum L.
Epipactis atrorubens (Hoffm.) B e s s e r
Epipactis helleborine (L.) C r a n t z
Eragrostis minor H o s t
Eriophorum angustifolium H o n c k.
Eriophorum vaginatum L.
Erodium cicutarium (L.) L' H é r.
Erophila verna (L.) C h e v a l l
Eupatorium cannabinum L.
Euphrasia rostkoviana H a y n e
Euphrasia stricta J. P. W o l f f
Fallopia convolvulus (L.) Á. L ö v e
Fallopia dumetorum (L.) H o l u b
Festuca arundinacea S c h r e b.
Festuca gigantea (L.) V i l l.
Festuca ovina L.
Festuca pratensis H u d s.
Festuca rubra L.
Filaginella uliginosa (L.) O p i t z
Filipendula ulmaria (L.) M a x i m.
Fragaria moschata (D u c h e s n e) W e s t o n
Fragaria vesca L.
Fumaria officinalis L.
Fumaria vaillantii L o i s e l.
Appendix. (Continued)
Poaceae
Apiaceae
Poaceae
Brassicaceae
Caryophyllaceae
Scrophulariaceae
Scrophulariaceae
Poaceae
Dipsacaceae
Poaceae
Boraginaceae
Cyperaceae
Poaceae
Onagraceae
Onagraceae
Onagraceae
Orchidaceae
Orchidaceae
Poaceae
Cyperaceae
Cyperaceae
Geraniaceae
Brassicaceae
Asteraceae
Scrophulariaceae
Scrophulariaceae
Polygonaceae
Polygonaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Asteraceae
Rosaceae
Rosaceae
Rosaceae
Fumariaceae
Fumariaceae
Family
caryopsis
mericarp
caryopsis
seed
seed
seed
seed
caryopsis
achene
caryopsis
nutlet
achene
caryopsis
seed
seed
seed
seed
seed
caryopsis
achene
achene
mericarp
seed
achene
seed
seed
achene
achene
caryopsis
caryopsis
caryopsis
caryopsis
caryopsis
achene
follicle
achene
achene
seed
seed
Diaspore
Reproductive
capacity
of population
34 648
168 025
162 199
426 042
217 359
376 605
1 002 454
270 460
384 264
145 242
115 915
73 667
13 715
3 857 071
1 343 250
2 478 960
15 333 333
13 679 278
920 875
40 191
14 203
44 525
3 836 667
120 597
577 250
1 096 600
12 962
15 816
116 325
37 843
178 025
125 631
119 777
10 308 221
135 460
69 750
49 967
27 040
17 725
Diaspore
weight
(mg)
1.074
1.003
0.350
0.140
0.131
0.140
0.099
0.405
3.050
0.800
2.740
–
1.860
0.625
0.108
0.045
–
0.002
0.093
0.555
0.590
2.547
0.013
0.238
0.117
0.202
3.845
3.160
2.395
2.433
–
1.467
0.645
0.007
0.500
–
0.330
0.960
0.840
Number
per life
PC
MC
PC
MC
PC
PC
MC
MC
MC
MC
MC
PC
PC
PC
PC
PC
PC
PC
MC
PC
PC
MC
MC
PC
MC
MC
MC
MC
PC
PC
PC
PC
PC
MC
PC
PC
PC
MC
MC
of flowering
Life
P
B
P
A
P
P
B
A
B
A
B
P
P
P
P
P
P
P
A
P
P
B
A
P
A
A
A
A
P
P
P
P
P
A
P
P
P
A
A
span
1
0
1
0
1
0
0
0
0
0
0
2
2
2
2
1
0
0
0
2
2
0
0
1
0
0
0
0
1
2
1
1
2
0
2
1
2
0
0
spread
Lateral
20
Name
Galeobdolon luteum H u d s. emend. H o l u b
Galeopsis bifida B o e n n.
Galeopsis pubescens B e s s e r
Galeopsis speciosa M i l l.
Galeopsis tetrahit L.
Galinsoga parviflora C a v.
Galium aparine L.
Galium mollugo L.
Galium odoratum (L.) S c o p.
Galium rotundifolium L.
Galium sylvaticum L.
Galium uliginosum L.
Galium verum L.
Genista tinctoria L.
Geranium dissectum L.
Geranium phaeum L.
Geranium pratense L.
Geranium pusillum B u r m. f.
Geranium robertianum L.
Geranium sanguineum L.
Geum urbanum L.
Glechoma hederacea L.
Glyceria fluitans (L.) R. B r.
Glyceria maxima (H a r t m.) H o l m b.
Gymnadenia conopsea (L.) R. B r.
Hedera helix L.
Helianthemum grandiflorum (S c o p.) D C.
Hepatica nobilis S c h r e b.
Heracleum mantegazzianum S o m m i e r et L e v i e r
Heracleum sphondylium L.
Herniaria glabra L.
Hieracium laevigatum W i l l d.
Hieracium murorum L.
Hieracium sabaudum L.
Hieracium umbellatum L.
Holcus lanatus L.
Holcus mollis L.
Homogyne alpina (L.) C a s s.
Hordeum jubatum L.
Appendix. (Continued)
Lamiaceae
Lamiaceae
Lamiaceae
Lamiaceae
Lamiaceae
Asteraceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Rubiaceae
Fabaceae
Geraniaceae
Geraniaceae
Geraniaceae
Geraniaceae
Geraniaceae
Geraniaceae
Rosaceae
Lamiaceae
Poaceae
Poaceae
Orchidaceae
Araliaceae
Cistaceae
Ranunculaceae
Apiaceae
Apiaceae
Caryophyllaceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Poaceae
Poaceae
Asteraceae
Poaceae
Family
nutlet
nutlet
nutlet
nutlet
nutlet
achene
mericarp
mericarp
mericarp
mericarp
mericarp
mericarp
mericarp
seed
mericarp
mericarp
mericarp
mericarp
mericarp
mericarp
achene
nutlet
caryopsis
caryopsis
seed
seed
seed
achene
mericarp
mericarp
seed
achene
achene
achene
achene
caryopsis
caryopsis
achene
caryopsis
Diaspore
Reproductive
capacity
of population
37 600
94 698
210 133
38 291
31 650
190 368
25 180
111 841
16 413
14 200
37 770
104 991
45 115
56 652
11 391
8 841
8 370
115 546
12 153
6 406
38 633
115 801
59 211
214 217
2 131 361
5 344
156 963
20 533
37 680
120 128
1 428 446
49 200
59 587
99 292
104 690
588 827
212 472
6 300
28 612
Diaspore
weight
(mg)
–
2.965
1.880
4.030
2.800
0.160
7.660
0.720
6.250
1.635
0.669
0.700
0.540
1.700
3.260
3.160
3.933
0.841
1.020
1.140
2.610
–
1.625
–
0.002
29.000
0.560
1.180
7.430
6.248
0.035
0.404
0.145
0.575
0.491
0.430
–
2.061
1.500
Number
per life
PC
MC
MC
MC
MC
MC
MC
PC
PC
PC
PC
PC
PC
PC
MC
PC
PC
MC
MC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
MC
PC
PC
PC
PC
PC
PC
PC
MC
of flowering
Life
P
A
A
A
A
A
A
P
P
P
P
P
P
P
A
P
P
A
A
P
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
P
P
P
A
span
2
0
0
0
0
0
0
1
1
1
2
1
1
1
0
1
1
0
0
2
1
2
2
2
0
0
1
0
0
1
0
1
0
2
1
1
2
1
0
spread
Lateral
21
Name
Hordeum murinum H u d s.
Hyosciamus niger L.
Hypericum perforatum L.
Hypochaeris radicata L.
Impatiens glandulifera R o y l e
Impatiens noli–tangere L.
Impatiens parviflora D C.
Iris pseudacorus L.
Iris pumila L.
Isopyrum thalictroides L.
Iva xanthiifolia N u t t.
Jacea pratensis L a m.
Jasione montana L.
Juncus articulatus L.
Juncus bufonius L.
Juncus conglomeratus L.
Juncus effusus L.
Juncus squarrosus L.
Juncus tenuis W i l l d.
Knautia arvensis (L.) C o u l t.
Knautia maxima (O p i z) O r t m a n n
Koeleria macranta (L e d e b.) S c h u l t.
Koeleria pyramidata (L a m.) P. B e a u v.
Lactuca serriola L.
Lamium album L.
Lamium amplexicaule L.
Lamium maculatum L.
Lamium purpureum L.
Lapsana communis L.
Lathraea squamaria L.
Lathyrus pratensis L.
Lathyrus sylvestris L.
Lathyrus vernus (L.) B e r n h.
Lembotropis nigricans (L.) G r i s e b.
Leontodon autumnalis L.
Leontodon hispidus L.
Leonurus cardiaca L.
Lepidium campestre (L.) R. B r.
Lepidium ruderale L.
Appendix. (Continued)
Poaceae
Solanaceae
Hypericaceae
Asteraceae
Balsaminaceae
Balsaminaceae
Balsaminaceae
Iridaceae
Iridaceae
Ranunculaceae
Asteraceae
Asteraceae
Campanulaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Dipsacaceae
Dipsacaceae
Poaceae
Poaceae
Asteraceae
Lamiaceae
Lamiaceae
Lamiaceae
Lamiaceae
Asteraceae
Scrophulariaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Asteraceae
Asteraceae
Lamiaceae
Brassicaceae
Brassicaceae
Family
caryopsis
seed
seed
achene
seed
seed
seed
seed
seed
seed
achene
achene
seed
seed
seed
seed
seed
seed
seed
achene
achene
caryopsis
caryopsis
achene
nutlet
nutlet
nutlet
nutlet
achene
seed
seed
seed
seed
seed
achene
achene
nutlet
seed
seed
Diaspore
Reproductive
capacity
of population
52 692
388 316
1 282 800
59 746
7 978
8 458
28 932
7 307
6 553
13 017
310 131
29 787
4 017 000
1 212 965
5 476 148
4 213 583
10 319 712
1 955 881
5 538 677
25 239
75 965
49 580
224 556
182 713
53 107
75 435
100 012
26 835
204 931
675 060
7 851
5 732
32 100
3 770
59 550
36 956
69 732
110 134
339 349
Diaspore
weight
(mg)
–
0.600
0.062
1.124
3.095
5.550
5.300
43.950
3.200
1.305
0.812
1.620
0.022
0.080
0.010
0.013
0.011
0.018
0.007
6.170
2.575
–
–
0.493
1.154
0.680
1.600
0.754
0.566
0.113
11.650
18.800
–
23.517
0.720
1.222
0.780
2.200
0.199
Number
per life
MC
MC
PC
PC
MC
MC
MC
PC
PC
PC
MC
PC
MC
PC
MC
PC
PC
PC
PC
PC
PC
PC
PC
MC
PC
MC
PC
MC
PC
PC
PC
PC
PC
PC
PC
PC
PC
MC
MC
of flowering
Life
A
B
P
P
A
A
A
P
P
P
A
P
B
P
A
P
P
P
P
P
P
P
P
B
P
A
P
A
P
P
P
P
P
P
P
P
P
A
B
span
0
0
1
0
0
0
0
1
1
1
0
1
0
1
0
1
1
1
1
0
1
1
1
0
2
0
1
0
0
0
1
1
1
1
0
1
2
0
0
spread
Lateral
22
Name
Leucanthemum ircutianum D C.
Leucojum vernum L.
Libanotis pyrenaica (L.) B o u r g.
Lilium martagon L.
Linaria genistifolia (L.) M i l l.
Linaria vulgaris M i l l.
Linum usitatissimum L.
Lithospermum arvense L.
Lolium multiflorum L a m.
Lolium perenne L.
Lotus corniculatus L.
Lunaria rediviva L.
Lupinus polyphyllus L i n d l.
Luzula campestris (L.) D C.
Luzula luzuloides (L a m.) D a n d y
Luzula multiflora (E h r h.) L e j.
Luzula pilosa (L.) W i l l d.
Luzula sylvatica (H u d s.) G a u d i n
Lychnis flos–cuculi L.
Lycopsis arvensis L.
Lycopus europaeus (L.)
Lysimachia nummularia L.
Lysimachia vulgaris L.
Lythrum salicaria L.
Maianthemum bifolium (L.) F. W. S c h m i d t
Malva neglecta W a l l r.
Matricaria discoidea D C.
Matricaria recutitaL.
Medicago lupulina L.
Medicago sativa L.
Melampyrum nemorosum L.
Melampyrum pratense L.
Melandrium latifolia P o i r.
Melica nutans L.
Melica transsilvanica S c h u r.
Melilotus albus M e d i k.
Melilotus officinalis (L.) P a l l.
Mentha arvensis L.
Mentha suaveolens E h r h.
Appendix. (Continued)
Asteraceae
Amaryllaceae
Apiaceae
Liliaceae
Scrophulariaceae
Scrophulariaceae
Linaceae
Boraginaceae
Poaceae
Poaceae
Fabaceae
Brassicaceae
Fabaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Caryophyllaceae
Boraginaceae
Lamiaceae
Primulaceae
Primulaceae
Lythraceae
Liliaceae
Malvaceae
Asteraceae
Asteraceae
Fabaceae
Fabaceae
Scrophulariaceae
Scrophulariaceae
Caryophyllaceae
Poaceae
Poaceae
Fabaceae
Fabaceae
Lamiaceae
Lamiaceae
Family
achene
seed
mericarp
seed
seed
seed
seed
nutlet
caryopsis
caryopsis
seed
seed
seed
seed
seed
seed
seed
seed
seed
nutlet
nutlet
seed
seed
seed
seed
mericarp
achene
achene
seed
seed
seed
seed
seed
caryopsis
caryopsis
seed
seed
nutlet
nutlet
Diaspore
Reproductive
capacity
of population
45 886
5 211
103 281
46 825
742 676
187 667
100 000
36 537
16 833
63 192
40 962
9 228
13 950
172 875
153 200
183 067
56 239
32 703
715 538
33 982
87 753
8 157
126 667
1 085 505
8 883
50 483
704 933
871 870
177 543
91 145
14 532
9 388
301 569
30 000
48 852
48 168
46 917
250 505
56 512
Diaspore
weight
(mg)
0.430
5.660
1.100
116.800
0.134
0.113
2.090
3.205
2.160
0.677
–
19.500
17.600
0.138
0.346
0.580
0.544
0.600
0.080
5.235
0.239
–
0.246
0.074
5.780
1.323
0.097
0.068
1.450
1.815
8.449
4.570
0.549
1.722
0.073
–
2.000
0.116
0.035
Number
per life
PC
PC
MC
PC
PC
PC
MC
MC
MC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
MC
PC
PC
PC
PC
PC
MC
MC
MC
MC
PC
MC
MC
PC
PC
PC
MC
MC
PC
PC
of flowering
Life
P
P
B
P
P
P
A
A
B
P
P
P
P
P
P
P
P
P
P
A
P
P
P
P
P
B
A
A
B
P
A
A
P
P
P
B
B
P
P
span
1
0
1
0
1
1
0
0
1
2
1
2
1
1
2
1
2
1
1
0
1
1
1
1
0
2
0
0
1
2
0
0
1
1
1
0
1
2
2
spread
Lateral
23
Name
Menyathes trifoliata L.
Merculialis perennis L.
Microrrhinum minus (L.) F o u r r.
Milium effusum L.
Mimulus guttatus D C.
Molinia caerulea (L.) M o e n c h
Moneses uniflora (L.) A. G r a y
Mycelis muralis (L.) D u m o r t.
Myosotis arvensis (L.) H i l l
Myosotis laxiflora R c h b.
Myosotis nemorosa B e s s e r
Myosotis sparsiflora M i k a n ex P o h l
Myosoton aquaticum (L.) M o e n c h
Nardus stricta L.
Neslia paniculata (L.) D e s v.
Odontites vulgaris M o e n c h
Oenanthe aquatica (L.) P o i r.
Oenothera biennis L.
Omalotheca sylvatica (L.) S c h. B i p. et F. W. S c h u l t z
Omphalodes scorpioides (H a e n k e) S c h r a n k
Onobrychis viciifolia S c o p.
Onopordum acanthium L.
Origanum vulgare L.
Orthantha lutea (L.) A. K e r n. ex W e t t s t.
Oxalis acetosella L.
Oxalis stricta L.
Oxyccocus palustris P e r s.
Papaver dubium L.
Papaver rhoeas L.
Pastinaca sativa L.
Pedicularis sylvatica L.
Pedicularis verticillata L.
Persicaria hydropiper (L.) S p a c h
Persicaria maculata (Rafin.) G r a y
Petasites hybridus (L.) P. G a e r t n., B. M e y et S c h e r b.
Peucedanum palustre (L.) M o e n c h
Phalaroides arundinacea (L.) R a u s c h e r t
Phleum pratense L.
Phragmites australis (C a v.) T r i n.
Appendix. (Continued)
Menyanthaceae
Euphorbiaceae
Scrophulariaceae
Poaceae
Scrophulariaceae
Poaceae
Pyrolaceae
Asteraceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Caryophyllaceae
Poaceae
Brassicaceae
Scrophulariaceae
Apiaceae
Oenotheraceae
Asteraceae
Boraginaceae
Fabaceae
Asteraceae
Lamiaceae
Scrophulariaceae
Oxalidaceae
Oxalidaceae
Ericaceae
Papaveraceae
Papaveraceae
Apiaceae
Scrophulariaceae
Scrophulariaceae
Polygonaceae
Polygonaceae
Asteraceae
Apiaceae
Poaceae
Poaceae
Poaceae
Family
seed
seed
seed
caryopsis
seed
caryopsis
seed
achene
nutlet
nutlet
nutlet
nutlet
seed
caryopsis
seed
seed
mericarp
seed
achene
nutlet
seed
achene
nutlet
seed
seed
seed
seed
seed
seed
mericarp
seed
seed
achene
achene
achene
mericarp
caryopsis
caryopsis
caryopsis
Diaspore
Reproductive
capacity
of population
1 626
1 726
948 603
63 794
1 041 627
68 879
1 155 440
44 287
116 197
51 257
33 859
49 477
692 357
13 122
214 913
952 098
47 100
588 804
1 004 711
39 867
10 617
12 751
217 340
165 255
4 179
96 800
5 317
877 825
683 950
141 172
41 490
27 440
44 640
5 970
10 839
102 608
29 829
45 790
63 200
Diaspore
weight
(mg)
–
3.215
0.083
0.510
0.017
0.039
0.001
0.218
0.275
0.269
0.407
0.820
0.076
0.510
1.480
0.147
1.070
0.190
0.043
0.195
5.920
10.770
0.057
0.202
1.050
0.166
0.870
0.077
0.095
1.735
0.922
1.730
1.900
2.680
0.355
2.600
0.719
0.568
1.575
Number
per life
PC
PC
MC
PC
PC
PC
PC
PC
MC
PC
PC
MC
PC
PC
MC
MC
MC
MC
PC
MC
PC
MC
PC
MC
PC
MC
PC
MC
MC
MC
PC
PC
MC
PC
PC
PC
PC
PC
PC
of flowering
Life
P
P
A
P
P
P
P
P
A
P
P
A
P
P
A
A
B
B
P
B
P
B
P
A
P
A
P
A
A
B
P
P
A
P
P
P
P
P
P
span
2
2
0
1
2
1
1
1
0
1
0
0
1
0
0
0
1
1
0
0
0
1
1
0
1
0
2
0
0
0
1
1
1
1
2
1
2
1
2
spread
Lateral
24
Name
Phyteuma nigrum S c h m.
Pimpinella major (L.) H u d s.
Pimpinella saxifraga L.
Plantago lanceolata L.
Plantago major L.
Poa annua L.
Poa compressa L.
Poa nemoralis L.
Poa pratensis L.
Polygonatum odoratum (M i l l.) D r u c e
Polygonum aviculare L.
Polygonum lapathifolia (L.) G r a y
Portulaca oleracea L.
Potamogeton pectinatus L.
Potentilla argentea L.
Potentilla erecta (L.) R d e u s c h.
Potentilla recta L.
Potentilla reptans L.
Potentilla supina L.
Potentilla tabernaemontani A s c h.
Prenanthes purpurea L.
Primula elatior (L.) H i l l .
Primula veris L.
Prunella vulgaris L.
Pseudofumaria lutea (L.) M e d i k.
Pseudolysimachion spicatum (L.) O p i z
Puccinellia distans (J a c q.) P a r l.
Pulmonaria officinalis L.
Pulsatilla grandis W e n d e r.
Pyrethrum corymbosum (L.) S c o p.
Pyrola minor L.
Ranunculus acris L.
Ranunculus auricomus L.
Ranunculus flammula L.
Ranunculus sceleratus L.
Raphanus raphanistrum L.
Reseda lutea L.
Reseda luteola L.
Rhinanthus minor L.
Appendix. (Continued)
Campanulaceae
Apiaceae
Apiaceae
Plantaginaceae
Plantaginaceae
Poaceae
Poaceae
Poaceae
Poaceae
Liliaceae
Polygonaceae
Polygonaceae
Portulacaceae
Potamogetonaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Asteraceae
Primulaceae
Primulaceae
Lamiaceae
Fumariaceae
Scrophulariaceae
Poaceae
Boraginaceae
Ranunculaceae
Asteraceae
Pyrolaceae
Ranunculaceae
Ranunculaceae
Ranunculaceae
Ranunculaceae
Brassicaceae
Resedaceae
Resedaceae
Scrophulariaceae
Family
seed
mericarp
mericarp
seed
seed
caryopsis
caryopsis
caryopsis
caryopsis
seed
achene
achene
seed
achene
achene
achene
achene
achene
achene
achene
achene
seed
seed
nutlet
seed
seed
caryopsis
nutlet
achene
achene
seed
achene
achene
achene
achene
seed
seed
seed
seed
Diaspore
Reproductive
capacity
of population
162 585
76 437
145 816
160 997
850 567
91 230
222 988
113 116
136 086
9 300
120 368
37 591
995 307
276
1 053 960
54 771
107 026
26 172
191 666
177 833
43 093
50 400
72 800
73 691
30 015
255 778
222 519
17 403
8 400
922 782
9 400 000
33 800
6 212
202 380
262 977
27 175
420 667
176 878
40 936
Diaspore
weight
(mg)
0.085
2.350
0.782
1.715
0.216
0.152
0.140
0.046
0.220
–
0.542
7.525
0.087
4.920
0.078
0.245
0.105
0.233
0.065
0.615
1.167
0.683
–
0.549
1.110
0.071
0.244
48.000
2.390
0.135
0.001
0.518
–
0.490
0.073
3.750
0.510
0.280
2.165
Number
per life
PC
PC
PC
PC
PC
MC
PC
PC
PC
PC
MC
MC
MC
PC
PC
PC
PC
PC
MC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
MC
MC
MC
MC
MC
of flowering
Life
P
P
P
P
P
A
P
P
P
P
A
A
A
P
P
P
P
P
B
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
A
A
B
B
A
span
1
1
1
0
0
0
1
2
1
1
0
0
0
2
1
2
0
1
1
0
2
0
1
1
0
0
0
0
0
0
1
0
0
2
0
0
1
1
0
spread
Lateral
25
Name
Rhinanthus pulcher G ü n t h e r et S c h u m m e l ex O p i z
Rhinanthus serotinus (S c h ö n h.) O b o r n y
Rorippa palustris (L.) B e s s e r
Rorippa sylvestris (L.) B e s s e r
Rumex crispus L.
Rumex maritimus L.
Rumex obtusifolius L.
Sagina procumbens L.
Salvia pratensis L.
Sanguisorba minor S c o p.
Sanguisorba officinalis L.
Sanicula europaea L.
Saponaria officinalis L.
Saxifraga granulata L.
Scirpus sylvaticus L.
Scorzonera humilis L.
Scrophularia nodosa L.
Scutellaria galericulata L.
Sedum acre L.
Selinum carvifolia (L.) L.
Senecio erraticus B e r t o l.
Senecio ovatus (P. G a e r t n., B. M e y et S c h e r b.) W i l l d.
Senecio rivularis (W. K.) D C.
Senecio sylvaticus L.
Senecio viscosus L.
Senecio vulgaris L.
Setaria pumila (P o i r.) R o e m. et S c h u l t.
Silaum silaus (L.) S c h i n z et T h e l l.
Silene dioica (L.) C l a i r v.
Silene nutans L.
Sinapis alba L.
Sinapis arvensis L.
Sisymbrium altissimum L.
Sisymbrium loeselii L.
Sisymbrium officinale (L.) S c o p.
Solanum dulcamara L.
Solanum nigrum L.
Solanum vilosum (L.) M i l l.
Soldanella montana W i l l d.
Appendix. (Continued)
Scrophulariaceae
Scrophulariaceae
Brassicaceae
Brassicaceae
Polygonaceae
Polygonaceae
Polygonaceae
Caryophyllaceae
Lamiaceae
Rosaceae
Rosaceae
Apiaceae
Caryophyllaceae
Saxifragaceae
Cyperaceae
Asteraceae
Scrophulariaceae
Lamiaceae
Crassulaceae
Apiaceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Poaceae
Apiaceae
Caryophyllaceae
Caryophyllaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Solanaceae
Solanaceae
Solanaceae
Primulaceae
Family
seed
seed
seed
seed
achene
achene
achene
seed
nutlet
achene
achene
mericarp
seed
seed
achene
achene
seed
nutlet
seed
mericarp
achene
achene
achene
achene
achene
achene
caryopsis
mericarp
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
Diaspore
Reproductive
capacity
of population
22 748
44 247
396 175
1 469 342
59 207
41 739
37 574
6 185 083
33 459
10 255
84 622
28 978
57 324
1 334 667
693 015
2 947
435 474
17 867
618 366
264 367
57 083
48 946
27 990
92 467
86 844
334 264
121 574
186 017
108 957
103 500
33 088
37 746
981 874
749 903
348 055
30 637
286 794
61 352
26 667
Diaspore
weight
(mg)
1.971
1.285
0.019
0.035
1.300
0.107
0.746
0.007
1.147
2.500
1.715
5.000
1.899
0.019
0.075
2.140
0.062
0.516
0.022
1.915
0.255
0.619
0.523
0.690
0.437
0.196
1.336
2.280
0.081
0.174
4.665
2.145
0.204
0.086
0.263
1.970
1.179
0.660
0.730
Number
per life
MC
MC
MC
PC
PC
MC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
MC
PC
PC
MC
MC
MC
MC
PC
PC
PC
MC
MC
MC
MC
MC
PC
MC
MC
PC
of flowering
Life
A
A
B
P
P
A
P
P
P
P
P
P
P
P
P
P
P
P
P
P
B
P
P
A
A
A
A
P
P
P
A
A
A
A
A
P
A
A
P
span
0
0
1
1
0
0
0
0
1
0
1
1
1
0
2
0
0
2
0
0
0
2
2
0
0
0
0
1
1
1
0
0
0
0
0
2
0
0
0
spread
Lateral
26
Name
Solidago gigantea A i t o n
Solidago virgaurea L.
Sonchus arvensis L.
Sonchus oleraceus L.
Sparganium erectum L.
Spergula arvensis L.
Spergularia rubra (L.) J. P r e s l et C. P r e s l
Stachys palustris L.
Stachys sylvatica L.
Stellaria graminea L.
Stellaria holostea L.
Steris viscaria (L.) R a f i n.
Symphytum tuberosum L.
Tanacetum vulgare L.
Taraxacum sect. Ruderalia K i r s c h n e r et al.
Teucrium chamaedrys L.
Thalictrum lucidum L.
Thlaspi arvense L.
Thlaspi perfoliatum L.
Thymus pulegioides L.
Tofieldia calyculata (L.) W a h l e n b.
Torilis japonica (Houtt.) D C.
Tragopogon pratensis L.
Trientalis europaea L.
Trifolium alpestre L.
Trifolium arvense L.
Trifolium aureum P o l l i c h
Trifolium campestre S c h r e b.
Trifolium dubium S i b t h.
Trifolium medium G r u f b.
Trifolium repens L.
Trifolium spadiceum L.
Tripleurospermum perforatum (M é r a t) M. L a í n z
Trisetum flavescens (L.) P. B e a u v.
Tussilago farfara L.
Typha latifolia L.
Tythymalus cyparissias (L.) S c o p.
Tythymalus esula (L.) H i l l .
Tythymalus helioscopia (L.) H i l l .
Appendix. (Continued)
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Sparganiaceae
Caryophyllaceae
Caryophyllaceae
Lamiaceae
Lamiaceae
Caryophyllaceae
Caryophyllaceae
Caryophyllaceae
Boraginaceae
Asteraceae
Asteraceae
Lamiaceae
Ranunculaceae
Brassicaceae
Brassicaceae
Lamiaceae
Liliaceae
Apiaceae
Asteraceae
Primulaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Asteraceae
Poaceae
Asteraceae
Typhaceae
Euphorbiaceae
Euphorbiaceae
Euphorbiaceae
Family
achene
achene
achene
achene
seed
seed
seed
nutlet
nutlet
seed
seed
seed
nutlet
achene
achene
nutlet
achene
seed
seed
nutlet
seed
mericarp
achene
seed
seed
seed
seed
seed
seed
seed
seed
seed
achene
caryopsis
achene
follicle
seed
seed
seed
Diaspore
Reproductive
capacity
of population
899 445
150 300
92 883
250 916
10 627
352 675
6 299 800
302 589
27 498
125 880
40 034
375 558
7 636
1 211 203
41 333
24 367
38 092
74 604
51 428
722 854
401 084
51 917
10 556
14 874
24 083
302 145
244 413
168 984
104 589
22 539
122 568
258 733
788 883
160 273
27 128
18 780 889
51 630
51 194
64 567
Diaspore
weight
(mg)
–
0.415
1.719
0.322
20.440
2.080
0.056
1.000
1.181
0.230
2.200
0.063
11.500
0.304
0.410
1.650
–
0.553
0.266
0.117
0.039
1.600
9.215
0.430
0.667
0.385
–
0.460
0.537
2.200
0.390
–
0.193
–
0.330
0.060
–
2.550
0.933
Number
per life
PC
PC
PC
MC
PC
MC
MC
PC
PC
PC
PC
PC
PC
PC
PC
PC
PC
MC
MC
PC
PC
MC
PC
PC
PC
MC
MC
MC
MC
PC
PC
MC
MC
PC
PC
PC
PC
PC
MC
of flowering
Life
P
P
P
A
P
A
A
P
P
P
P
P
P
P
P
P
P
A
A
P
P
B
P
P
P
A
A
A
A
P
P
A
A
P
P
P
P
P
A
span
2
1
1
0
2
0
0
2
1
1
1
1
1
2
0
1
1
0
0
1
1
0
1
0
1
0
0
0
0
1
2
0
0
1
2
2
2
1
0
spread
Lateral
27
Name
Urtica dioica L.
Urtica urens L.
Vaccinium myrtillus L.
Valeriana dioica L.
Valerianella locusta (L.) L a t e r r.
Verbascum chaixii V i l l.
Verbascum nigrum L.
Verbascum thapsus L.
Veronica arvensis L.
Veronica chamaedrys L.
Veronica hederifolia L.
Veronica montana L.
Veronica officinalis L.
Veronica persica P o i r.
Veronica scutellata L.
Veronica serpyllifolia L.
Veronica triphyllos L.
Vicia cracca L.
Vicia hirsuta (L.) G r a y
Vicia pannonica C r a n t z.
Vicia sepium L.
Vicia tetrasperma (L.) S c h r e b.
Vicia villosa R o t h
Vincetoxicum hirundinaria M e d i k.
Viola arvensis M u r r a y
Viola canina L. emend. R c h b.
Viola palustris L.
Viola reichenbachiana J o r d. ex B o r e a u
Appendix. (Continued)
Urticaceae
Urticaceae
Ericaceae
Valerianaceae
Valerianaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Fabaceae
Asplepiadaceae
Violaceae
Violaceae
Violaceae
Violaceae
Family
seed
seed
seed
achene
achene
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
seed
Diaspore
Reproductive
capacity
of population
35 674
107 820
5 237
3 180
44 000
142 439
486 399
2 489 337
504 581
293 431
1 366 667
38 400
183 082
136 911
999 935
1 671 528
228 633
41 867
26 333
25 933
11 781
27 850
23 373
120 670
103 800
28 442
18 130
15 000
Diaspore
weight
(mg)
0.167
0.479
–
0.400
0.275
0.114
1.783
0.097
0.517
0.187
0.518
–
0.150
0.535
0.012
0.026
0.137
11.680
6.600
19.800
18.300
2.760
38.900
2.490
0.175
0.648
0.290
1.190
Number
per life
PC
MC
PC
PC
MC
MC
MC
MC
MC
PC
MC
PC
PC
MC
PC
PC
MC
PC
MC
MC
PC
MC
MC
PC
MC
PC
PC
PC
of flowering
Life
P
A
P
P
A
B
B
B
A
P
A
P
P
A
P
P
A
P
A
A
P
A
A
P
A
P
P
P
span
2
0
1
2
0
0
0
0
0
1
0
1
1
0
1
1
0
1
0
0
1
0
0
1
0
1
1
1
spread
Lateral