DIASPORES POTENTIAL OR REAL POWER OF WILD PLANTS? LIFE CYCLE

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: sera@usbe.cas.cz 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 flow- 7

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 veg- 8

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 n Mean Media n SD Reproductive capacity of population (1 000/m 2 ) Diaspore weight (mg) Number of flowering Monocarps a 175 540 116 1 469 Polycarps b 319 500 59 1 861 Life span Annuals a 131 595 120 1 637 Biennials and Monocarpic perennials ab 44 377 99 772 Perennials b 319 500 59 1 861 Lateral spread Without a 224 659 107 1 930 < 0.30 m per year b 176 384 68 1 225 0.31 m per year b 92 408 76 2 010 Number of flowering Monocarps a 169 2.03 0.68 4.42 Polycarps a 286 2.88 0.65 8.98 Life span Annuals a 126 1.79 0.58 4.13 Biennials and Monocarpic perennials a 43 2.72 1.07 5.17 Perennials a 286 2.88 0.65 8.98 Lateral spread Without a 219 3.10 0.68 9.80 < 0.30 m per year a 157 2.21 0.70 4.95 0.31 m per year a 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 strate- 9

7,0 8 6,5 7 Reproductive capacity of population 6,0 5,5 5,0 4,5 4,0 3,5 Reproductive capacity of population 6 5 4 3 3,0 2,5 Monocarps Polycarps 2 Annuals Biennials Perennials Number of flowering Life span 8 7 Reproductive capacity of population 6 5 4 3 2 0 1 2 Lateral spread 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

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. Number of flowering Life span Lateral spread Diaspore weight Reproductive capacity of population Number of flowering 1.000 Life span ****0.982 1.000 Lateral spread ****0.607 ****0.630 1.000 Diaspore weight -0.006 0.007-0.024 1.000 Reproductive capacity of population *** - 0.197 **** - 0.200 **- 0.171 **- 0.760 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 in a dependence on specific life conditions (e.g. A rgen, Willson, 1994). 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. References A rgen, J., Willson, M.F., 1994: Cost of seed production in the herbs Geranium maculatum and G. sylvaticum: an experimental field study. Oikos, 70, p. 35 42. Baker, H.G., 1972: Seed weight in relation to environmental conditions in California. Ecology, 53, p. 997 1010. Begon, M., Harper, J.L., Townsend, C.R., 1990: Ecology: Individuals, Populations and Communities. Blackwell Scientific Publications, London, 949 pp. Boyd, M., Silvertown, J., Tucker, C., 1990: Population of heterostyle and homostyle Primula vulgaris: growth, survival and reproduction in field populations. J. Ecol., 78, p. 799 813. Cavers, P.B., Steele, M.G., 1984: Patterns of change in seed weight over time on individual plants. Am. Nat., 124, p. 324 335. Colosi, J.C., Cavers, P.B., 1984: Pollination affects percent biomass allocated to reproduction in Silene vulgaris bladder champion. Am. Nat., 124, p. 299 306. Cornelissen, J.H.C., Lavorel, S., Garnier, E., Díaz, S., Buchmann, N., Gurvich, D.E., Reich, P.B., ter Steege, H., Morgan, H.D., van der Heijden, M.G.A, Pausas, J.G., Poorter, H., 2003: A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust. J. Bot., 51, p. 335 380. Crawley, M.J., 1987: What makes a community invisible? In Gray, A.J., Crawley, M.J., Edwards, P.J. (eds): Colonization, Succession and Stability. Blackwell Scientic, Oxford, p. 429 543. Downs, C., McQuilkin, W.E., 1944: Seed production of Southern Appalachian oaks. J. Forestry, 42, p. 786 790. Fenner, M., 1985: Seed Ecology. Chapman and Hall, London, 151 pp. Francland, B., 1975: Phytochrome control of seed germination in relation to the light environment. In Smith, H. (ed.): Light and Plant Development. Butterworth, London, p. 477 502. 12

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Wardlaw, I.F., Dunstone, R.L., 1984: Effect of temperature on seed development in jujube (Simmondsia chinensis (L i n k) S c h n e d e r). 1. Dry matter changes. Aust. J. Agric. Res., 35, p. 685 691. Westoby, M., Leischman, M., Janice, L., 1996: Comparative ecology of seed size and dispersal. Philos. Trans. R. Soc. Lond. B, 351, p. 1309 1318. Wulff, R.D., 1986: Seed size variation in Desmodium paniculatum. I. Factors affecting seed size. J. Ecol., 74, p. 87 97. Šerá B.: Diaspory potenciální nebo reálná síla divokých rostlin? Životní cyklus. Received 25. 7. 2005 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

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. Achillea millefolium L. Asteraceae achene 0.033 101 882 PC P 1 Achillea ptarmica L. Asteraceae achene 66 400 PC P 1 Aconitum variegatum L. Ranunculaceae seed 1.907 35 122 PC P 1 Actaea spicata L. Ranunculaceae seed 5.895 1 164 PC P 0 Aethusa cynapium L. Apiaceae mericarp 1.600 126 314 MC A 0 Agrimonia procera W a l l r. Rosaceae achene 2.000 9 265 PC P 1 Agrostis canina L. Poaceae caryopsis 0.047 385 148 PC P 2 Agrostis capillaris L. Poaceae caryopsis 0.056 491 234 PC P 2 Agrostis stolonifera L. Poaceae caryopsis 0.045 62 463 PC P 2 Ajuga genevensis L. Lamiaceae nutlet 124 933 PC P 2 Ajuga reptans L. Lamiaceae nutlet 0.730 53 753 PC P 2 Alchemilla monticola O p i z Rosaceae achene 0.221 55 413 PC P 1 Alisma plantago aquatica L. Alismataceae achene 0.250 330 444 PC P 2 Alliaria petiolata (M. B i e b.) C a v a r a et G r a n d e Brassicaceae seed 2.090 138 508 MC A 0 Allium ursinum L. Liliaceae seed 3.365 4 751 PC P 0 Alopecurus pratensis L. Poaceae caryopsis 0.520 90 309 PC P 1 Alyssum alyssoides (L.) L. Brassicaceae seed 0.381 103 913 MC A 0 Amaranthus retroflexus L. Amaranthaceae seed 0.409 138 123 MC A 0 Anagallis arvensis L. Primulaceae seed 0.256 212 971 MC A 0 Anemone nemorosa L. Ranunculaceae achene 1.283 60 182 PC P 2 Angelica sylvestris L. Apiaceae mericarp 2.500 15 643 MC B 1 Anthemis arvensis L. Asteraceae achene 0.130 114 100 MC A 0 Anthoxanthum odoratum L. Poaceae caryopsis 0.286 211 895 PC P 1 Anthriscus sylvestris (L.) H o f f m. Apiaceae mericarp 2.033 62 448 PC P 0 Apera spica venti (L.) P. B e a u v. Poaceae caryopsis 0.141 250 801 MC A 0 Aphanes arvensis L. Rosaceae achene 0.172 242 205 MC A 0 Arabidopsis thaliana (L.) H e y n h. Brassicaceae seed 0.012 4 448 014 MC A 0 Arabis glabra (L.) B e r n h. Brassicaceae seed 0.014 1 483 500 MC A 0 Arabis hirsuta (L.) S c o p. Brassicaceae seed 0.074 1 103 333 MC B 0 Arctium lappa L. Asteraceae achene 13.378 15 167 MC B 0 Arctium tomentosum M i l l. Asteraceae achene 11.130 29 117 MC B 0 Arenaria serpyllifolia L. Caryophyllaceae seed 0.025 1 256 927 MC A 0 Arnica montana L. Asteraceae achene 0.955 67 304 PC P 0 Arrhenatherum elatius (L.) J. P r e s l et C. P r e s l Poaceae caryopsis 2.130 25 151 PC P 1 Artemisia vulgaris L. Asteraceae achene 0.090 177 548 PC P 1 15

Aruncus vulgaris R a f i n. Rosaceae follicle 0.090 32 459 PC P 2 Asarum europaeum L. Aristolochiacea seed 3.740 8 467 PC P 1 Astragalus glycyphyllos L. Fabaceae seed 3.970 7 221 PC P 1 Astrantia major L. Apiaceae mericarp 1.800 41 293 PC P 0 Atriplex patula L. Chenopodiaceae achene 7.600 67 945 MC A 0 Atriplex sagittata B o r k h. Chenopodiaceae achene 1.000 40 959 MC A 0 Atropa bella dona L. Solanaceae seed 1.030 32 613 PC P 0 Avenella flexuosa (L.) P a r l. Poaceae caryopsis 0.266 81 857 PC P 1 Ballota nigra L. Lamiaceae nutlet 1.675 91 670 PC P 0 Barbarea vulgaris R. B r. Brassicaceae seed 0.600 212 286 MC B 0 Batrachium aquatile (L.) D u m o r t Ranunculaceae achene 88 204 MC A 2 Bellis perennis L. Asteraceae achene 0.082 67 407 PC P 0 Berteroa incana (L.) D C. Brassicaceae seed 0.270 139 388 MC A 0 Betonica officinalis L. Lamiaceae nutlet 1.159 109 926 PC P 0 Bidens cernua L. Asteraceae achene 0.907 53 041 MC A 0 Bidens frondosa L. Asteraceae achene 2.056 53 308 MC A 0 Bidens tripartita L. Asteraceae achene 2.100 231 371 MC A 0 Brachypodium pinnatum (L.) P. B e a u v. Poaceae caryopsis 65 000 PC P 2 Brachypodium sylvaticum (H u d s.) P. B e a u v. Poaceae caryopsis 2.730 11 131 PC P 1 Brasica napus L. Brassicaceae seed 180 472 MC A 0 Briza media L. Poaceae caryopsis 0.523 30 956 PC P 1 Bromus erectus H u d s. Poaceae caryopsis 120 964 PC P 1 Bromus hordeaceus L. Poaceae caryopsis 2.100 63 610 MC A 0 Bromus inermis L e y s s Poaceae caryopsis 27 077 PC P 1 Bromus sterilis L. Poaceae caryopsis 3.900 67 835 MC A 0 Bromus tectorum L. Poaceae caryopsis 2.000 105 400 MC A 0 Bryonia alba L. Cucurbitaceae seed 11.800 1 619 PC P 0 Butomus umbellatus L. Butomaceae seed 0.200 124 796 PC P 1 Calamagrostis arundinacea (L.) R o t h Poaceae caryopsis 0.380 15 887 PC P 2 Calamagrostis canescens (W e b e r) R o t h Poaceae caryopsis 0.200 295 597 PC P 2 Calamagrostis epigejos (L.) R o t h Poaceae caryopsis 0.256 258 327 PC P 2 Calamagrostis villosa (C h a i x ex V i l l.) J. F. G m e l Poaceae caryopsis 0.817 34 102 PC P 2 Callitriche palustris agg. Callitrichaceae seed 0.023 148 290 MC A 0 Calluna vulgaris (L.) H u l l Ericaceae seed 0.060 761 890 PC P 1 Caltha palustris L. Ranunculaceae seed 0.663 245 794 PC P 0 Calystegia sepium (L.) R. B r. Convolvulaceae seed 22.870 969 PC P 0 Camelina microcarpa A n d r z. Brassicaceae seed 0.180 373 133 PC P 0 Campanula patula L. Campanulaceae seed 0.020 2 375 000 MC B 0 Campanula persicifolia L. Campanulaceae seed 0.080 375 000 PC P 0 16

Campanula rapunculoides L. Campanulaceae seed 0.220 175 000 PC P 1 Campanula rotundifolia L. Campanulaceae seed 0.030 2 143 146 PC P 1 Campanula trachelium L. Campanulaceae seed 0.143 268 837 PC P 2 Capsella bursa pastoris (L.) M e d i k. Brassicaceae seed 0.113 1 965 033 MC A 0 Cardamine pratensis L. Brassicaceae seed 0.162 236 918 PC P 0 Cardaminopsis halleri (L.) H a y e k Brassicaceae seed 3 521 393 PC P 1 Carduus crispus L. Asteraceae achene 1.902 21 200 MC B 1 Carduus nutans L. Asteraceae achene 1.968 3 498 MC B 1 Carduus personata (L.) J a c q. Asteraceae achene 2.540 6 113 PC P 1 Carex brizoides L. Cyperaceae achene 0.260 44 620 PC P 2 Carex canescens L. Cyperaceae achene 0.153 55 143 PC P 2 Carex elata A l l. Cyperaceae achene 0.835 22 615 PC P 1 Carex flacca S c h r e b. Cyperaceae achene 0.886 12 080 PC P 2 Carex gracilis C u r t i s. Cyperaceae achene 0.450 27 863 PC P 2 Carex hirta L. Cyperaceae achene 0.910 38 232 PC P 2 Carex humilis L e y s s. Cyperaceae achene 1.780 14 000 PC P 1 Carex nigra (L.) R e i c h a r d Cyperaceae achene 1.643 118 629 PC P 2 Carex ovalis G o o d e n. Cyperaceae achene 0.274 48 912 PC P 1 Carex pairaei F. W. S c h u l t z Cyperaceae achene 0.080 40 776 PC P 1 Carex pallescens L. Cyperaceae achene 1.230 64 256 PC P 1 Carex panicea L. Cyperaceae achene 0.985 37 269 PC P 2 Carex pilosa S c o p. Cyperaceae achene 1.000 9 761 PC P 1 Carex praecox S c h r e b. Cyperaceae achene 52 274 PC P 1 Carex riparia C u r t i s Cyperaceae achene 23 261 PC P 2 Carex rostrata S t o k e s ex W i t h. Cyperaceae achene 0.563 26 575 PC P 2 Carex spicata H u d s. Cyperaceae achene 1.700 27 900 PC P 1 Carex vesicaria L. Cyperaceae achene 0.543 29 776 PC P 2 Carex vulpina L. Cyperaceae achene 1.430 17 967 PC P 1 Carlina acaulis L. Asteraceae achene 3.460 29 556 PC P 0 Carum carvi L. Apiaceae mericarp 1.050 95 191 MC B 0 Centaurium erythraea R a f n Gentianaceae seed 0.004 12 252 500 MC A 0 Cerastium arvense L. Caryophyllaceae seed 0.116 560 841 PC P 1 Cerastium holosteoides F r. Caryophyllaceae seed 0.091 545 067 PC P 0 Cerastium pumilum C u r t. Caryophyllaceae seed 0.035 3 061 602 MC A 0 Chaerophyllum aromaticum L. Apiaceae mericarp 11.200 79 401 PC P 1 Chaerophyllum aureum L. Apiaceae mericarp 7.300 77 067 PC P 0 Chaerophyllum hirsutum L. Apiaceae mericarp 2.100 15 184 PC P 1 Chaerophyllum temulum L. Apiaceae mericarp 10.400 86 914 MC B 0 Chamerion angustifolium (L.) H o l u b Onagraceae seed 0.104 2 930 167 PC P 2 17

Chelidonium majus L. Papaveraceae seed 0.697 286 799 PC P 1 Chenopodium album L. Chenopodiaceae achene 0.656 312 589 MC A 0 Chenopodium bonus henricus L. Chenopodiaceae achene 1.990 23 442 PC P 1 Chenopodium glaucum L. Chenopodiaceae achene 0.102 110 556 MC A 0 Chenopodium hybridum L. Chenopodiaceae achene 1.296 228 120 MC A 0 Chenopodium pedunculare B e r t o l. Chenopodiaceae achene 0.698 206 018 MC A 0 Chenopodium polyspermum L. Chenopodiaceae achene 0.239 264 321 MC A 0 Chenopodium rubrum L. Chenopodiaceae achene 0.114 222 192 MC A 0 Chenopodium strictum R o t h Chenopodiaceae achene 0.330 65 468 MC A 0 Chenopodium suecicum M u r r Chenopodiaceae achene 0.620 79 928 MC A 0 Cichorium intybus L. Asteraceae achene 0.738 76 096 PC P 0 Cicuta virosa L. Apiaceae mericarp 2.200 30 500 PC P 1 Circaea luteciana L. Onagraceae seed 0.550 5 806 PC P 2 Cirsium arvense (L.) S c o p. Asteraceae achene 1.485 32 776 PC P 2 Cirsium heterophyllum (L.) H i l l Asteraceae achene 2.330 5 750 PC P 2 Cirsium oleraceum (L.) S c o p. Asteraceae achene 3.640 13 985 PC P 2 Cirsium palustre (L.) S c o p. Asteraceae achene 1.202 59 469 MC B 1 Cirsium vulgare (S a v i) T e n. Asteraceae achene 4.627 53 070 MC B 2 Clinopodium vulgare L. Lamiaceae nutlet 0.452 216 818 PC P 1 Colchicum autumnale L. Liliaceae seed 7.900 86 767 PC P 0 Conium maculatum L. Apiaceae mericarp 1.000 92 388 MC B 0 Consolida regalis G r a y Ranunculaceae seed 0.830 197 647 MC A 0 Convallaria majalis L. Liliaceae seed 26.160 2 469 PC P 2 Convolvulus arvensis L. Convolvulaceae seed 11.370 909 PC P 2 Conyza canadensis (L.) C r o n g u i s t Asteraceae achene 0.048 800 476 MC A 0 Coronilla varia L. Fabaceae seed 0.810 129 900 PC P 2 Corydalis cava (L.) S c h w e i g g. et K ö r t e Fumariaceae seed 1.225 17 664 PC P 0 Corydalis solida (L.) C l a i r v. Fumariaceae seed 5.250 13 183 PC P 0 Crepis biennis L. Asteraceae achene 0.982 74 896 MC B 2 Cruciata glabra (L.) E h r e n d. Rubiaceae mericarp 243 185 PC P 1 Cuscuta europaea L. Cuscutaceae seed 0.600 190 748 MC A 0 Cyanus segetum H i l l Asteraceae achene 2.870 25 166 MC A 0 Cymbalaria muralis P. G a e r t n., B. M e y. et S c h e r b. Scrophulariaceae seed 0.128 59 560 PC P 2 Cynodon dactylon (L.) P e r s. Poaceae caryopsis 55 550 PC P 2 Cynoglossum officinale L. Boraginaceae nutlet 28.700 14 453 MC B 0 Cynosurus cristatus L. Poaceae caryopsis 2.800 96 897 PC P 1 Dactylis glomerata L. Poaceae caryopsis 0.963 138 622 PC P 1 Dactylorhiza fuchsii (D r u c e) S o ó Orchidaceae seed 0.001 4 074 563 PC P 0 Dactylorhiza majalis (R c h b.) P. F. H u n t et S u m m e r h Orchidaceae seed 0.002 4 720 600 PC P 0 18

Danthonia decumbens (L.) D C. Poaceae caryopsis 1.074 34 648 PC P 1 Daucus carota L. Apiaceae mericarp 1.003 168 025 MC B 0 Deschampsia cespitosa (L.) P. B e a u v. Poaceae caryopsis 0.350 162 199 PC P 1 Descurainia sophia (L.) W e b b ex P r a n t l Brassicaceae seed 0.140 426 042 MC A 0 Dianthus deltoides L. Caryophyllaceae seed 0.131 217 359 PC P 1 Digitalis grandiflora M i l l. Scrophulariaceae seed 0.140 376 605 PC P 0 Digitalis purpurea L. Scrophulariaceae seed 0.099 1 002 454 MC B 0 Digitaria sanguinalis (L.) S c o p. Poaceae caryopsis 0.405 270 460 MC A 0 Dipsacus fullonum L. Dipsacaceae achene 3.050 384 264 MC B 0 Echinochloa crus galli (L.) P. B e a u v. Poaceae caryopsis 0.800 145 242 MC A 0 Echium vulgare L. Boraginaceae nutlet 2.740 115 915 MC B 0 Eleocharis palustris (L.) R o e m. et S c h u l t. Cyperaceae achene 73 667 PC P 2 Elytrigia repens (L.) D e s v. Poaceae caryopsis 1.860 13 715 PC P 2 Epilobium ciliatum R a f. Onagraceae seed 0.625 3 857 071 PC P 2 Epilobium hirsutum L. Onagraceae seed 0.108 1 343 250 PC P 2 Epilobium montanum L. Onagraceae seed 0.045 2 478 960 PC P 1 Epipactis atrorubens (Hoffm.) B e s s e r Orchidaceae seed 15 333 333 PC P 0 Epipactis helleborine (L.) C r a n t z Orchidaceae seed 0.002 13 679 278 PC P 0 Eragrostis minor H o s t Poaceae caryopsis 0.093 920 875 MC A 0 Eriophorum angustifolium H o n c k. Cyperaceae achene 0.555 40 191 PC P 2 Eriophorum vaginatum L. Cyperaceae achene 0.590 14 203 PC P 2 Erodium cicutarium (L.) L' H é r. Geraniaceae mericarp 2.547 44 525 MC B 0 Erophila verna (L.) C h e v a l l Brassicaceae seed 0.013 3 836 667 MC A 0 Eupatorium cannabinum L. Asteraceae achene 0.238 120 597 PC P 1 Euphrasia rostkoviana H a y n e Scrophulariaceae seed 0.117 577 250 MC A 0 Euphrasia stricta J. P. W o l f f Scrophulariaceae seed 0.202 1 096 600 MC A 0 Fallopia convolvulus (L.) Á. L ö v e Polygonaceae achene 3.845 12 962 MC A 0 Fallopia dumetorum (L.) H o l u b Polygonaceae achene 3.160 15 816 MC A 0 Festuca arundinacea S c h r e b. Poaceae caryopsis 2.395 116 325 PC P 1 Festuca gigantea (L.) V i l l. Poaceae caryopsis 2.433 37 843 PC P 2 Festuca ovina L. Poaceae caryopsis 178 025 PC P 1 Festuca pratensis H u d s. Poaceae caryopsis 1.467 125 631 PC P 1 Festuca rubra L. Poaceae caryopsis 0.645 119 777 PC P 2 Filaginella uliginosa (L.) O p i t z Asteraceae achene 0.007 10 308 221 MC A 0 Filipendula ulmaria (L.) M a x i m. Rosaceae follicle 0.500 135 460 PC P 2 Fragaria moschata (D u c h e s n e) W e s t o n Rosaceae achene 69 750 PC P 1 Fragaria vesca L. Rosaceae achene 0.330 49 967 PC P 2 Fumaria officinalis L. Fumariaceae seed 0.960 27 040 MC A 0 Fumaria vaillantii L o i s e l. Fumariaceae seed 0.840 17 725 MC A 0 19

Galeobdolon luteum H u d s. emend. H o l u b Lamiaceae nutlet 37 600 PC P 2 Galeopsis bifida B o e n n. Lamiaceae nutlet 2.965 94 698 MC A 0 Galeopsis pubescens B e s s e r Lamiaceae nutlet 1.880 210 133 MC A 0 Galeopsis speciosa M i l l. Lamiaceae nutlet 4.030 38 291 MC A 0 Galeopsis tetrahit L. Lamiaceae nutlet 2.800 31 650 MC A 0 Galinsoga parviflora C a v. Asteraceae achene 0.160 190 368 MC A 0 Galium aparine L. Rubiaceae mericarp 7.660 25 180 MC A 0 Galium mollugo L. Rubiaceae mericarp 0.720 111 841 PC P 1 Galium odoratum (L.) S c o p. Rubiaceae mericarp 6.250 16 413 PC P 1 Galium rotundifolium L. Rubiaceae mericarp 1.635 14 200 PC P 1 Galium sylvaticum L. Rubiaceae mericarp 0.669 37 770 PC P 2 Galium uliginosum L. Rubiaceae mericarp 0.700 104 991 PC P 1 Galium verum L. Rubiaceae mericarp 0.540 45 115 PC P 1 Genista tinctoria L. Fabaceae seed 1.700 56 652 PC P 1 Geranium dissectum L. Geraniaceae mericarp 3.260 11 391 MC A 0 Geranium phaeum L. Geraniaceae mericarp 3.160 8 841 PC P 1 Geranium pratense L. Geraniaceae mericarp 3.933 8 370 PC P 1 Geranium pusillum B u r m. f. Geraniaceae mericarp 0.841 115 546 MC A 0 Geranium robertianum L. Geraniaceae mericarp 1.020 12 153 MC A 0 Geranium sanguineum L. Geraniaceae mericarp 1.140 6 406 PC P 2 Geum urbanum L. Rosaceae achene 2.610 38 633 PC P 1 Glechoma hederacea L. Lamiaceae nutlet 115 801 PC P 2 Glyceria fluitans (L.) R. B r. Poaceae caryopsis 1.625 59 211 PC P 2 Glyceria maxima (H a r t m.) H o l m b. Poaceae caryopsis 214 217 PC P 2 Gymnadenia conopsea (L.) R. B r. Orchidaceae seed 0.002 2 131 361 PC P 0 Hedera helix L. Araliaceae seed 29.000 5 344 PC P 0 Helianthemum grandiflorum (S c o p.) D C. Cistaceae seed 0.560 156 963 PC P 1 Hepatica nobilis S c h r e b. Ranunculaceae achene 1.180 20 533 PC P 0 Heracleum mantegazzianum S o m m i e r et L e v i e r Apiaceae mericarp 7.430 37 680 PC P 0 Heracleum sphondylium L. Apiaceae mericarp 6.248 120 128 PC P 1 Herniaria glabra L. Caryophyllaceae seed 0.035 1 428 446 MC A 0 Hieracium laevigatum W i l l d. Asteraceae achene 0.404 49 200 PC P 1 Hieracium murorum L. Asteraceae achene 0.145 59 587 PC P 0 Hieracium sabaudum L. Asteraceae achene 0.575 99 292 PC P 2 Hieracium umbellatum L. Asteraceae achene 0.491 104 690 PC P 1 Holcus lanatus L. Poaceae caryopsis 0.430 588 827 PC P 1 Holcus mollis L. Poaceae caryopsis 212 472 PC P 2 Homogyne alpina (L.) C a s s. Asteraceae achene 2.061 6 300 PC P 1 Hordeum jubatum L. Poaceae caryopsis 1.500 28 612 MC A 0 20

Hordeum murinum H u d s. Poaceae caryopsis 52 692 MC A 0 Hyosciamus niger L. Solanaceae seed 0.600 388 316 MC B 0 Hypericum perforatum L. Hypericaceae seed 0.062 1 282 800 PC P 1 Hypochaeris radicata L. Asteraceae achene 1.124 59 746 PC P 0 Impatiens glandulifera R o y l e Balsaminaceae seed 3.095 7 978 MC A 0 Impatiens noli tangere L. Balsaminaceae seed 5.550 8 458 MC A 0 Impatiens parviflora D C. Balsaminaceae seed 5.300 28 932 MC A 0 Iris pseudacorus L. Iridaceae seed 43.950 7 307 PC P 1 Iris pumila L. Iridaceae seed 3.200 6 553 PC P 1 Isopyrum thalictroides L. Ranunculaceae seed 1.305 13 017 PC P 1 Iva xanthiifolia N u t t. Asteraceae achene 0.812 310 131 MC A 0 Jacea pratensis L a m. Asteraceae achene 1.620 29 787 PC P 1 Jasione montana L. Campanulaceae seed 0.022 4 017 000 MC B 0 Juncus articulatus L. Juncaceae seed 0.080 1 212 965 PC P 1 Juncus bufonius L. Juncaceae seed 0.010 5 476 148 MC A 0 Juncus conglomeratus L. Juncaceae seed 0.013 4 213 583 PC P 1 Juncus effusus L. Juncaceae seed 0.011 10 319 712 PC P 1 Juncus squarrosus L. Juncaceae seed 0.018 1 955 881 PC P 1 Juncus tenuis W i l l d. Juncaceae seed 0.007 5 538 677 PC P 1 Knautia arvensis (L.) C o u l t. Dipsacaceae achene 6.170 25 239 PC P 0 Knautia maxima (O p i z) O r t m a n n Dipsacaceae achene 2.575 75 965 PC P 1 Koeleria macranta (L e d e b.) S c h u l t. Poaceae caryopsis 49 580 PC P 1 Koeleria pyramidata (L a m.) P. B e a u v. Poaceae caryopsis 224 556 PC P 1 Lactuca serriola L. Asteraceae achene 0.493 182 713 MC B 0 Lamium album L. Lamiaceae nutlet 1.154 53 107 PC P 2 Lamium amplexicaule L. Lamiaceae nutlet 0.680 75 435 MC A 0 Lamium maculatum L. Lamiaceae nutlet 1.600 100 012 PC P 1 Lamium purpureum L. Lamiaceae nutlet 0.754 26 835 MC A 0 Lapsana communis L. Asteraceae achene 0.566 204 931 PC P 0 Lathraea squamaria L. Scrophulariaceae seed 0.113 675 060 PC P 0 Lathyrus pratensis L. Fabaceae seed 11.650 7 851 PC P 1 Lathyrus sylvestris L. Fabaceae seed 18.800 5 732 PC P 1 Lathyrus vernus (L.) B e r n h. Fabaceae seed 32 100 PC P 1 Lembotropis nigricans (L.) G r i s e b. Fabaceae seed 23.517 3 770 PC P 1 Leontodon autumnalis L. Asteraceae achene 0.720 59 550 PC P 0 Leontodon hispidus L. Asteraceae achene 1.222 36 956 PC P 1 Leonurus cardiaca L. Lamiaceae nutlet 0.780 69 732 PC P 2 Lepidium campestre (L.) R. B r. Brassicaceae seed 2.200 110 134 MC A 0 Lepidium ruderale L. Brassicaceae seed 0.199 339 349 MC B 0 21

Leucanthemum ircutianum D C. Asteraceae achene 0.430 45 886 PC P 1 Leucojum vernum L. Amaryllaceae seed 5.660 5 211 PC P 0 Libanotis pyrenaica (L.) B o u r g. Apiaceae mericarp 1.100 103 281 MC B 1 Lilium martagon L. Liliaceae seed 116.800 46 825 PC P 0 Linaria genistifolia (L.) M i l l. Scrophulariaceae seed 0.134 742 676 PC P 1 Linaria vulgaris M i l l. Scrophulariaceae seed 0.113 187 667 PC P 1 Linum usitatissimum L. Linaceae seed 2.090 100 000 MC A 0 Lithospermum arvense L. Boraginaceae nutlet 3.205 36 537 MC A 0 Lolium multiflorum L a m. Poaceae caryopsis 2.160 16 833 MC B 1 Lolium perenne L. Poaceae caryopsis 0.677 63 192 PC P 2 Lotus corniculatus L. Fabaceae seed 40 962 PC P 1 Lunaria rediviva L. Brassicaceae seed 19.500 9 228 PC P 2 Lupinus polyphyllus L i n d l. Fabaceae seed 17.600 13 950 PC P 1 Luzula campestris (L.) D C. Juncaceae seed 0.138 172 875 PC P 1 Luzula luzuloides (L a m.) D a n d y Juncaceae seed 0.346 153 200 PC P 2 Luzula multiflora (E h r h.) L e j. Juncaceae seed 0.580 183 067 PC P 1 Luzula pilosa (L.) W i l l d. Juncaceae seed 0.544 56 239 PC P 2 Luzula sylvatica (H u d s.) G a u d i n Juncaceae seed 0.600 32 703 PC P 1 Lychnis flos cuculi L. Caryophyllaceae seed 0.080 715 538 PC P 1 Lycopsis arvensis L. Boraginaceae nutlet 5.235 33 982 MC A 0 Lycopus europaeus (L.) Lamiaceae nutlet 0.239 87 753 PC P 1 Lysimachia nummularia L. Primulaceae seed 8 157 PC P 1 Lysimachia vulgaris L. Primulaceae seed 0.246 126 667 PC P 1 Lythrum salicaria L. Lythraceae seed 0.074 1 085 505 PC P 1 Maianthemum bifolium (L.) F. W. S c h m i d t Liliaceae seed 5.780 8 883 PC P 0 Malva neglecta W a l l r. Malvaceae mericarp 1.323 50 483 MC B 2 Matricaria discoidea D C. Asteraceae achene 0.097 704 933 MC A 0 Matricaria recutital. Asteraceae achene 0.068 871 870 MC A 0 Medicago lupulina L. Fabaceae seed 1.450 177 543 MC B 1 Medicago sativa L. Fabaceae seed 1.815 91 145 PC P 2 Melampyrum nemorosum L. Scrophulariaceae seed 8.449 14 532 MC A 0 Melampyrum pratense L. Scrophulariaceae seed 4.570 9 388 MC A 0 Melandrium latifolia P o i r. Caryophyllaceae seed 0.549 301 569 PC P 1 Melica nutans L. Poaceae caryopsis 1.722 30 000 PC P 1 Melica transsilvanica S c h u r. Poaceae caryopsis 0.073 48 852 PC P 1 Melilotus albus M e d i k. Fabaceae seed 48 168 MC B 0 Melilotus officinalis (L.) P a l l. Fabaceae seed 2.000 46 917 MC B 1 Mentha arvensis L. Lamiaceae nutlet 0.116 250 505 PC P 2 Mentha suaveolens E h r h. Lamiaceae nutlet 0.035 56 512 PC P 2 22

Menyathes trifoliata L. Menyanthaceae seed 1 626 PC P 2 Merculialis perennis L. Euphorbiaceae seed 3.215 1 726 PC P 2 Microrrhinum minus (L.) F o u r r. Scrophulariaceae seed 0.083 948 603 MC A 0 Milium effusum L. Poaceae caryopsis 0.510 63 794 PC P 1 Mimulus guttatus D C. Scrophulariaceae seed 0.017 1 041 627 PC P 2 Molinia caerulea (L.) M o e n c h Poaceae caryopsis 0.039 68 879 PC P 1 Moneses uniflora (L.) A. G r a y Pyrolaceae seed 0.001 1 155 440 PC P 1 Mycelis muralis (L.) D u m o r t. Asteraceae achene 0.218 44 287 PC P 1 Myosotis arvensis (L.) H i l l Boraginaceae nutlet 0.275 116 197 MC A 0 Myosotis laxiflora R c h b. Boraginaceae nutlet 0.269 51 257 PC P 1 Myosotis nemorosa B e s s e r Boraginaceae nutlet 0.407 33 859 PC P 0 Myosotis sparsiflora M i k a n ex P o h l Boraginaceae nutlet 0.820 49 477 MC A 0 Myosoton aquaticum (L.) M o e n c h Caryophyllaceae seed 0.076 692 357 PC P 1 Nardus stricta L. Poaceae caryopsis 0.510 13 122 PC P 0 Neslia paniculata (L.) D e s v. Brassicaceae seed 1.480 214 913 MC A 0 Odontites vulgaris M o e n c h Scrophulariaceae seed 0.147 952 098 MC A 0 Oenanthe aquatica (L.) P o i r. Apiaceae mericarp 1.070 47 100 MC B 1 Oenothera biennis L. Oenotheraceae seed 0.190 588 804 MC B 1 Omalotheca sylvatica (L.) S c h. B i p. et F. W. S c h u l t z Asteraceae achene 0.043 1 004 711 PC P 0 Omphalodes scorpioides (H a e n k e) S c h r a n k Boraginaceae nutlet 0.195 39 867 MC B 0 Onobrychis viciifolia S c o p. Fabaceae seed 5.920 10 617 PC P 0 Onopordum acanthium L. Asteraceae achene 10.770 12 751 MC B 1 Origanum vulgare L. Lamiaceae nutlet 0.057 217 340 PC P 1 Orthantha lutea (L.) A. K e r n. ex W e t t s t. Scrophulariaceae seed 0.202 165 255 MC A 0 Oxalis acetosella L. Oxalidaceae seed 1.050 4 179 PC P 1 Oxalis stricta L. Oxalidaceae seed 0.166 96 800 MC A 0 Oxyccocus palustris P e r s. Ericaceae seed 0.870 5 317 PC P 2 Papaver dubium L. Papaveraceae seed 0.077 877 825 MC A 0 Papaver rhoeas L. Papaveraceae seed 0.095 683 950 MC A 0 Pastinaca sativa L. Apiaceae mericarp 1.735 141 172 MC B 0 Pedicularis sylvatica L. Scrophulariaceae seed 0.922 41 490 PC P 1 Pedicularis verticillata L. Scrophulariaceae seed 1.730 27 440 PC P 1 Persicaria hydropiper (L.) S p a c h Polygonaceae achene 1.900 44 640 MC A 1 Persicaria maculata (Rafin.) G r a y Polygonaceae achene 2.680 5 970 PC P 1 Petasites hybridus (L.) P. G a e r t n., B. M e y et S c h e r b. Asteraceae achene 0.355 10 839 PC P 2 Peucedanum palustre (L.) M o e n c h Apiaceae mericarp 2.600 102 608 PC P 1 Phalaroides arundinacea (L.) R a u s c h e r t Poaceae caryopsis 0.719 29 829 PC P 2 Phleum pratense L. Poaceae caryopsis 0.568 45 790 PC P 1 Phragmites australis (C a v.) T r i n. Poaceae caryopsis 1.575 63 200 PC P 2 23

Phyteuma nigrum S c h m. Campanulaceae seed 0.085 162 585 PC P 1 Pimpinella major (L.) H u d s. Apiaceae mericarp 2.350 76 437 PC P 1 Pimpinella saxifraga L. Apiaceae mericarp 0.782 145 816 PC P 1 Plantago lanceolata L. Plantaginaceae seed 1.715 160 997 PC P 0 Plantago major L. Plantaginaceae seed 0.216 850 567 PC P 0 Poa annua L. Poaceae caryopsis 0.152 91 230 MC A 0 Poa compressa L. Poaceae caryopsis 0.140 222 988 PC P 1 Poa nemoralis L. Poaceae caryopsis 0.046 113 116 PC P 2 Poa pratensis L. Poaceae caryopsis 0.220 136 086 PC P 1 Polygonatum odoratum (M i l l.) D r u c e Liliaceae seed 9 300 PC P 1 Polygonum aviculare L. Polygonaceae achene 0.542 120 368 MC A 0 Polygonum lapathifolia (L.) G r a y Polygonaceae achene 7.525 37 591 MC A 0 Portulaca oleracea L. Portulacaceae seed 0.087 995 307 MC A 0 Potamogeton pectinatus L. Potamogetonaceae achene 4.920 276 PC P 2 Potentilla argentea L. Rosaceae achene 0.078 1 053 960 PC P 1 Potentilla erecta (L.) R d e u s c h. Rosaceae achene 0.245 54 771 PC P 2 Potentilla recta L. Rosaceae achene 0.105 107 026 PC P 0 Potentilla reptans L. Rosaceae achene 0.233 26 172 PC P 1 Potentilla supina L. Rosaceae achene 0.065 191 666 MC B 1 Potentilla tabernaemontani A s c h. Rosaceae achene 0.615 177 833 PC P 0 Prenanthes purpurea L. Asteraceae achene 1.167 43 093 PC P 2 Primula elatior (L.) H i l l. Primulaceae seed 0.683 50 400 PC P 0 Primula veris L. Primulaceae seed 72 800 PC P 1 Prunella vulgaris L. Lamiaceae nutlet 0.549 73 691 PC P 1 Pseudofumaria lutea (L.) M e d i k. Fumariaceae seed 1.110 30 015 PC P 0 Pseudolysimachion spicatum (L.) O p i z Scrophulariaceae seed 0.071 255 778 PC P 0 Puccinellia distans (J a c q.) P a r l. Poaceae caryopsis 0.244 222 519 PC P 0 Pulmonaria officinalis L. Boraginaceae nutlet 48.000 17 403 PC P 0 Pulsatilla grandis W e n d e r. Ranunculaceae achene 2.390 8 400 PC P 0 Pyrethrum corymbosum (L.) S c o p. Asteraceae achene 0.135 922 782 PC P 0 Pyrola minor L. Pyrolaceae seed 0.001 9 400 000 PC P 1 Ranunculus acris L. Ranunculaceae achene 0.518 33 800 PC P 0 Ranunculus auricomus L. Ranunculaceae achene 6 212 PC P 0 Ranunculus flammula L. Ranunculaceae achene 0.490 202 380 PC P 2 Ranunculus sceleratus L. Ranunculaceae achene 0.073 262 977 MC A 0 Raphanus raphanistrum L. Brassicaceae seed 3.750 27 175 MC A 0 Reseda lutea L. Resedaceae seed 0.510 420 667 MC B 1 Reseda luteola L. Resedaceae seed 0.280 176 878 MC B 1 Rhinanthus minor L. Scrophulariaceae seed 2.165 40 936 MC A 0 24

Rhinanthus pulcher G ü n t h e r et S c h u m m e l ex O p i z Scrophulariaceae seed 1.971 22 748 MC A 0 Rhinanthus serotinus (S c h ö n h.) O b o r n y Scrophulariaceae seed 1.285 44 247 MC A 0 Rorippa palustris (L.) B e s s e r Brassicaceae seed 0.019 396 175 MC B 1 Rorippa sylvestris (L.) B e s s e r Brassicaceae seed 0.035 1 469 342 PC P 1 Rumex crispus L. Polygonaceae achene 1.300 59 207 PC P 0 Rumex maritimus L. Polygonaceae achene 0.107 41 739 MC A 0 Rumex obtusifolius L. Polygonaceae achene 0.746 37 574 PC P 0 Sagina procumbens L. Caryophyllaceae seed 0.007 6 185 083 PC P 0 Salvia pratensis L. Lamiaceae nutlet 1.147 33 459 PC P 1 Sanguisorba minor S c o p. Rosaceae achene 2.500 10 255 PC P 0 Sanguisorba officinalis L. Rosaceae achene 1.715 84 622 PC P 1 Sanicula europaea L. Apiaceae mericarp 5.000 28 978 PC P 1 Saponaria officinalis L. Caryophyllaceae seed 1.899 57 324 PC P 1 Saxifraga granulata L. Saxifragaceae seed 0.019 1 334 667 PC P 0 Scirpus sylvaticus L. Cyperaceae achene 0.075 693 015 PC P 2 Scorzonera humilis L. Asteraceae achene 2.140 2 947 PC P 0 Scrophularia nodosa L. Scrophulariaceae seed 0.062 435 474 PC P 0 Scutellaria galericulata L. Lamiaceae nutlet 0.516 17 867 PC P 2 Sedum acre L. Crassulaceae seed 0.022 618 366 PC P 0 Selinum carvifolia (L.) L. Apiaceae mericarp 1.915 264 367 PC P 0 Senecio erraticus B e r t o l. Asteraceae achene 0.255 57 083 MC B 0 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. Asteraceae achene 0.619 48 946 PC P 2 Senecio rivularis (W. K.) D C. Asteraceae achene 0.523 27 990 PC P 2 Senecio sylvaticus L. Asteraceae achene 0.690 92 467 MC A 0 Senecio viscosus L. Asteraceae achene 0.437 86 844 MC A 0 Senecio vulgaris L. Asteraceae achene 0.196 334 264 MC A 0 Setaria pumila (P o i r.) R o e m. et S c h u l t. Poaceae caryopsis 1.336 121 574 MC A 0 Silaum silaus (L.) S c h i n z et T h e l l. Apiaceae mericarp 2.280 186 017 PC P 1 Silene dioica (L.) C l a i r v. Caryophyllaceae seed 0.081 108 957 PC P 1 Silene nutans L. Caryophyllaceae seed 0.174 103 500 PC P 1 Sinapis alba L. Brassicaceae seed 4.665 33 088 MC A 0 Sinapis arvensis L. Brassicaceae seed 2.145 37 746 MC A 0 Sisymbrium altissimum L. Brassicaceae seed 0.204 981 874 MC A 0 Sisymbrium loeselii L. Brassicaceae seed 0.086 749 903 MC A 0 Sisymbrium officinale (L.) S c o p. Brassicaceae seed 0.263 348 055 MC A 0 Solanum dulcamara L. Solanaceae seed 1.970 30 637 PC P 2 Solanum nigrum L. Solanaceae seed 1.179 286 794 MC A 0 Solanum vilosum (L.) M i l l. Solanaceae seed 0.660 61 352 MC A 0 Soldanella montana W i l l d. Primulaceae seed 0.730 26 667 PC P 0 25

Solidago gigantea A i t o n Asteraceae achene 899 445 PC P 2 Solidago virgaurea L. Asteraceae achene 0.415 150 300 PC P 1 Sonchus arvensis L. Asteraceae achene 1.719 92 883 PC P 1 Sonchus oleraceus L. Asteraceae achene 0.322 250 916 MC A 0 Sparganium erectum L. Sparganiaceae seed 20.440 10 627 PC P 2 Spergula arvensis L. Caryophyllaceae seed 2.080 352 675 MC A 0 Spergularia rubra (L.) J. P r e s l et C. P r e s l Caryophyllaceae seed 0.056 6 299 800 MC A 0 Stachys palustris L. Lamiaceae nutlet 1.000 302 589 PC P 2 Stachys sylvatica L. Lamiaceae nutlet 1.181 27 498 PC P 1 Stellaria graminea L. Caryophyllaceae seed 0.230 125 880 PC P 1 Stellaria holostea L. Caryophyllaceae seed 2.200 40 034 PC P 1 Steris viscaria (L.) R a f i n. Caryophyllaceae seed 0.063 375 558 PC P 1 Symphytum tuberosum L. Boraginaceae nutlet 11.500 7 636 PC P 1 Tanacetum vulgare L. Asteraceae achene 0.304 1 211 203 PC P 2 Taraxacum sect. Ruderalia K i r s c h n e r et al. Asteraceae achene 0.410 41 333 PC P 0 Teucrium chamaedrys L. Lamiaceae nutlet 1.650 24 367 PC P 1 Thalictrum lucidum L. Ranunculaceae achene 38 092 PC P 1 Thlaspi arvense L. Brassicaceae seed 0.553 74 604 MC A 0 Thlaspi perfoliatum L. Brassicaceae seed 0.266 51 428 MC A 0 Thymus pulegioides L. Lamiaceae nutlet 0.117 722 854 PC P 1 Tofieldia calyculata (L.) W a h l e n b. Liliaceae seed 0.039 401 084 PC P 1 Torilis japonica (Houtt.) D C. Apiaceae mericarp 1.600 51 917 MC B 0 Tragopogon pratensis L. Asteraceae achene 9.215 10 556 PC P 1 Trientalis europaea L. Primulaceae seed 0.430 14 874 PC P 0 Trifolium alpestre L. Fabaceae seed 0.667 24 083 PC P 1 Trifolium arvense L. Fabaceae seed 0.385 302 145 MC A 0 Trifolium aureum P o l l i c h Fabaceae seed 244 413 MC A 0 Trifolium campestre S c h r e b. Fabaceae seed 0.460 168 984 MC A 0 Trifolium dubium S i b t h. Fabaceae seed 0.537 104 589 MC A 0 Trifolium medium G r u f b. Fabaceae seed 2.200 22 539 PC P 1 Trifolium repens L. Fabaceae seed 0.390 122 568 PC P 2 Trifolium spadiceum L. Fabaceae seed 258 733 MC A 0 Tripleurospermum perforatum (M é r a t) M. L a í n z Asteraceae achene 0.193 788 883 MC A 0 Trisetum flavescens (L.) P. B e a u v. Poaceae caryopsis 160 273 PC P 1 Tussilago farfara L. Asteraceae achene 0.330 27 128 PC P 2 Typha latifolia L. Typhaceae follicle 0.060 18 780 889 PC P 2 Tythymalus cyparissias (L.) S c o p. Euphorbiaceae seed 51 630 PC P 2 Tythymalus esula (L.) H i l l. Euphorbiaceae seed 2.550 51 194 PC P 1 Tythymalus helioscopia (L.) H i l l. Euphorbiaceae seed 0.933 64 567 MC A 0 26