DIASPORES POTENTIAL OR REAL POWER OF WILD PLANTS? LIFE CYCLE

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1 Ekológia (Bratislava) Vol. 24, Supplement 1/2005, p 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, České Budějovice, The Czech Republic, 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 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

2 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

3 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 Polycarps b Life span Annuals a Biennials and Monocarpic perennials ab Perennials b Lateral spread Without a < 0.30 m per year b m per year b Number of flowering Monocarps a Polycarps a Life span Annuals a Biennials and Monocarpic perennials a Perennials a Lateral spread Without a < 0.30 m per year a m per year a 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

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

5 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 < ***, P < ****. See the text. Number of flowering Life span Lateral spread Diaspore weight Reproductive capacity of population Number of flowering Life span **** Lateral spread ****0.607 **** Diaspore weight Reproductive capacity of population *** **** ** ** 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

6 (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. AV0Z 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 Baker, H.G., 1972: Seed weight in relation to environmental conditions in California. Ecology, 53, p 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 Cavers, P.B., Steele, M.G., 1984: Patterns of change in seed weight over time on individual plants. Am. Nat., 124, p Colosi, J.C., Cavers, P.B., 1984: Pollination affects percent biomass allocated to reproduction in Silene vulgaris bladder champion. Am. Nat., 124, p 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 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 Downs, C., McQuilkin, W.E., 1944: Seed production of Southern Appalachian oaks. J. Forestry, 42, p 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

7 Frank, D., Klotz, S., 1988: Biologisch ökologische Daten zur Flora der DDR. MLU Halle Wittenberg, 103 pp. Galen, C., Stanton, M.L., 1991: Consequences of emergence phenology for reproductive success in Ranunculus adoneus (Ranunculaceae). Am. J. Bot., 78, p Grime, J.P., 1977: Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. Am. Nat., 111, p Grime, J.P., Hillier, S.H., 1992: The contribution of seedling regeneration to the structure and dynamics of plant communities and larger units of landscape. In Fenner, M. (ed.): Seeds. The ecology of regeneration in plant communities, C.A.B. International, Wallingfound, p Grime, J.P., 2001: Plant Strategies, Vegetation Processes, and Ecosystem Properties. J. Wiley and Sons, Chichester, 415 pp. Harper, J.L., 1977: Population Biology of Plants. Academic Press, London, 892 pp. Hejný, S., Slavík, B. (eds), 1988: Flora of the Czech Socialistic Republic, 1 (in Czech). Academia, Praha, 557 pp. Hejný, S., Slavík, B. (eds), 1990: Flora of the Czech Republic, 2 (in Czech). Academia, Praha, 540 pp. Hejný, S., Slavík, B. (eds), 1992: Flora of the Czech Republic, 3 (in Czech). Academia, Praha, 542 pp. Hendrix, S.D., 1984: Variation in seed weight and its effects on germination in Pastinaca sativa L. (Umbelliferae). Am. J. Bot., 71, p Hodgson, J.G., Mackey, J.M.L., 1986: The ecological specialisation of dicotyledonous families within a local flora: some factors constraining optimalisation of seed size and their possible evolutionary significance. New Phytol., 104, p Jakrlová, J., 1985: Seed production of a Polygalo-Nardetum grassland. 1. Quantitative changes recorded in the course of years. Ekológia (ČSSR), 4, 2, p Kubát, K., Hrouda, L., Chrtek, J. jun., Kaplan, Z., Kirschner, J., Štěpánek, J. (eds), 2002: Key to the Flora of the Czech Republic (in Czech). Academia, Praha, 928 pp. Lhotská, M., Krippelová, T., Cigánová, K., 1987: How Reproduce and Disperse the Plants? (in Slovak). Obzor, Bratislava, 390 pp. Lloyd, D.G., 1987: Selection of offspring size at independence and other size-versus-number strategies. Am. Nat., 129, p Lovett Doust, J., 1980: A comparative study of life history and resource allocation in selected Umbelliferae. Biol. J. Linn. Soc., 13, p Pijl, L., 1982: Principles of Dispersal in Higher Plants. Springer Verlag, Berlin, Heidelberg, New York, 214 pp. Pyšek, P., Jarošík, V., Chytrý, M., Kropáč, Z., Tichý, L., Wild, J., 2005: Alien plants in temperate weed communities: prehistoric and recent invaders occupy different habitats. Ecology, 86, 3, p Rosenzweig, M.L., 1995: Species Diversity in Space and Time. Cambridge University Press, Cambridge, 136 pp. Rothmaler, W., Jäger, E., Schubert, R., Werner, K., 1987: Exkursionsflora für die Gebiete der DDR und der BRD. Atlas der Gefäßpflanzen. Volk und Wissen Volfseigener Verlag, Berlin, 752 pp. Salisbury, E.J., 1942: The Reproductive Capacity Per Plants. Bell and Sons, London, 244 pp. Silvertown, J.W., 1981: Seed size, lifespan and germination date as co-adapted features of plant life history. Am. Nat., 118, p Silvertown, J.W., Lovett Doust, J.L., 1993: Introduction to Plant Population Biology. Blackwell Scientific Publications, Oxford, 210 pp. Slavík, B. (ed.), 1995: Flora of the Czech Republic, 4 (in Czech). Academia, Praha, 529 pp. Slavík, B. (ed.), 1997: Flora of the Czech Republic, 5 (in Czech). Academia, Praha, 568 pp. Southwood, T.R.E., 1988: Tactics, strategies and templets. Oikos, 52, p Stanton, M.L., 1984: Developmental and genetic sources of seed weight variation in Raphanus raphanistrum L. (Brassicaceae). Am. J. Bot., 71, p Šerá, B., 1995: Reproductive capacity of the flowering plants (in Czech). Biol. Listy, 60, 2, p Šerá, B., Šerý, M., 2004: Relation between number and weight of seeds and reproductive strategies in herbaceous plants. Folia Geobot., 39, p Thompson, J.N., 1984: Variation among individual seed masses in Lomatium grayi (Umbelliferae) under controlled conditions: magnitude and partitioning of variance. Ecology, 65, p Waller, D.M., 1982: Factors influencing seed weight in Impatiens capensis (Balsaminaceae). Am. J. Bot., 69, p

8 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 Westoby, M., Leischman, M., Janice, L., 1996: Comparative ecology of seed size and dispersal. Philos. Trans. R. Soc. Lond. B, 351, p Wulff, R.D., 1986: Seed size variation in Desmodium paniculatum. I. Factors affecting seed size. J. Ecol., 74, p Šerá B.: Diaspory potenciální nebo reálná síla divokých rostlin? Životní cyklus. Received 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

9 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 PC P 1 Achillea ptarmica L. Asteraceae achene PC P 1 Aconitum variegatum L. Ranunculaceae seed PC P 1 Actaea spicata L. Ranunculaceae seed PC P 0 Aethusa cynapium L. Apiaceae mericarp MC A 0 Agrimonia procera W a l l r. Rosaceae achene PC P 1 Agrostis canina L. Poaceae caryopsis PC P 2 Agrostis capillaris L. Poaceae caryopsis PC P 2 Agrostis stolonifera L. Poaceae caryopsis PC P 2 Ajuga genevensis L. Lamiaceae nutlet PC P 2 Ajuga reptans L. Lamiaceae nutlet PC P 2 Alchemilla monticola O p i z Rosaceae achene PC P 1 Alisma plantago aquatica L. Alismataceae achene 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 MC A 0 Allium ursinum L. Liliaceae seed PC P 0 Alopecurus pratensis L. Poaceae caryopsis PC P 1 Alyssum alyssoides (L.) L. Brassicaceae seed MC A 0 Amaranthus retroflexus L. Amaranthaceae seed MC A 0 Anagallis arvensis L. Primulaceae seed MC A 0 Anemone nemorosa L. Ranunculaceae achene PC P 2 Angelica sylvestris L. Apiaceae mericarp MC B 1 Anthemis arvensis L. Asteraceae achene MC A 0 Anthoxanthum odoratum L. Poaceae caryopsis PC P 1 Anthriscus sylvestris (L.) H o f f m. Apiaceae mericarp PC P 0 Apera spica venti (L.) P. B e a u v. Poaceae caryopsis MC A 0 Aphanes arvensis L. Rosaceae achene MC A 0 Arabidopsis thaliana (L.) H e y n h. Brassicaceae seed MC A 0 Arabis glabra (L.) B e r n h. Brassicaceae seed MC A 0 Arabis hirsuta (L.) S c o p. Brassicaceae seed MC B 0 Arctium lappa L. Asteraceae achene MC B 0 Arctium tomentosum M i l l. Asteraceae achene MC B 0 Arenaria serpyllifolia L. Caryophyllaceae seed MC A 0 Arnica montana L. Asteraceae achene PC P 0 Arrhenatherum elatius (L.) J. P r e s l et C. P r e s l Poaceae caryopsis PC P 1 Artemisia vulgaris L. Asteraceae achene PC P 1 15

10 Aruncus vulgaris R a f i n. Rosaceae follicle PC P 2 Asarum europaeum L. Aristolochiacea seed PC P 1 Astragalus glycyphyllos L. Fabaceae seed PC P 1 Astrantia major L. Apiaceae mericarp PC P 0 Atriplex patula L. Chenopodiaceae achene MC A 0 Atriplex sagittata B o r k h. Chenopodiaceae achene MC A 0 Atropa bella dona L. Solanaceae seed PC P 0 Avenella flexuosa (L.) P a r l. Poaceae caryopsis PC P 1 Ballota nigra L. Lamiaceae nutlet PC P 0 Barbarea vulgaris R. B r. Brassicaceae seed MC B 0 Batrachium aquatile (L.) D u m o r t Ranunculaceae achene MC A 2 Bellis perennis L. Asteraceae achene PC P 0 Berteroa incana (L.) D C. Brassicaceae seed MC A 0 Betonica officinalis L. Lamiaceae nutlet PC P 0 Bidens cernua L. Asteraceae achene MC A 0 Bidens frondosa L. Asteraceae achene MC A 0 Bidens tripartita L. Asteraceae achene MC A 0 Brachypodium pinnatum (L.) P. B e a u v. Poaceae caryopsis PC P 2 Brachypodium sylvaticum (H u d s.) P. B e a u v. Poaceae caryopsis PC P 1 Brasica napus L. Brassicaceae seed MC A 0 Briza media L. Poaceae caryopsis PC P 1 Bromus erectus H u d s. Poaceae caryopsis PC P 1 Bromus hordeaceus L. Poaceae caryopsis MC A 0 Bromus inermis L e y s s Poaceae caryopsis PC P 1 Bromus sterilis L. Poaceae caryopsis MC A 0 Bromus tectorum L. Poaceae caryopsis MC A 0 Bryonia alba L. Cucurbitaceae seed PC P 0 Butomus umbellatus L. Butomaceae seed PC P 1 Calamagrostis arundinacea (L.) R o t h Poaceae caryopsis PC P 2 Calamagrostis canescens (W e b e r) R o t h Poaceae caryopsis PC P 2 Calamagrostis epigejos (L.) R o t h Poaceae caryopsis PC P 2 Calamagrostis villosa (C h a i x ex V i l l.) J. F. G m e l Poaceae caryopsis PC P 2 Callitriche palustris agg. Callitrichaceae seed MC A 0 Calluna vulgaris (L.) H u l l Ericaceae seed PC P 1 Caltha palustris L. Ranunculaceae seed PC P 0 Calystegia sepium (L.) R. B r. Convolvulaceae seed PC P 0 Camelina microcarpa A n d r z. Brassicaceae seed PC P 0 Campanula patula L. Campanulaceae seed MC B 0 Campanula persicifolia L. Campanulaceae seed PC P 0 16

11 Campanula rapunculoides L. Campanulaceae seed PC P 1 Campanula rotundifolia L. Campanulaceae seed PC P 1 Campanula trachelium L. Campanulaceae seed PC P 2 Capsella bursa pastoris (L.) M e d i k. Brassicaceae seed MC A 0 Cardamine pratensis L. Brassicaceae seed PC P 0 Cardaminopsis halleri (L.) H a y e k Brassicaceae seed PC P 1 Carduus crispus L. Asteraceae achene MC B 1 Carduus nutans L. Asteraceae achene MC B 1 Carduus personata (L.) J a c q. Asteraceae achene PC P 1 Carex brizoides L. Cyperaceae achene PC P 2 Carex canescens L. Cyperaceae achene PC P 2 Carex elata A l l. Cyperaceae achene PC P 1 Carex flacca S c h r e b. Cyperaceae achene PC P 2 Carex gracilis C u r t i s. Cyperaceae achene PC P 2 Carex hirta L. Cyperaceae achene PC P 2 Carex humilis L e y s s. Cyperaceae achene PC P 1 Carex nigra (L.) R e i c h a r d Cyperaceae achene PC P 2 Carex ovalis G o o d e n. Cyperaceae achene PC P 1 Carex pairaei F. W. S c h u l t z Cyperaceae achene PC P 1 Carex pallescens L. Cyperaceae achene PC P 1 Carex panicea L. Cyperaceae achene PC P 2 Carex pilosa S c o p. Cyperaceae achene PC P 1 Carex praecox S c h r e b. Cyperaceae achene PC P 1 Carex riparia C u r t i s Cyperaceae achene PC P 2 Carex rostrata S t o k e s ex W i t h. Cyperaceae achene PC P 2 Carex spicata H u d s. Cyperaceae achene PC P 1 Carex vesicaria L. Cyperaceae achene PC P 2 Carex vulpina L. Cyperaceae achene PC P 1 Carlina acaulis L. Asteraceae achene PC P 0 Carum carvi L. Apiaceae mericarp MC B 0 Centaurium erythraea R a f n Gentianaceae seed MC A 0 Cerastium arvense L. Caryophyllaceae seed PC P 1 Cerastium holosteoides F r. Caryophyllaceae seed PC P 0 Cerastium pumilum C u r t. Caryophyllaceae seed MC A 0 Chaerophyllum aromaticum L. Apiaceae mericarp PC P 1 Chaerophyllum aureum L. Apiaceae mericarp PC P 0 Chaerophyllum hirsutum L. Apiaceae mericarp PC P 1 Chaerophyllum temulum L. Apiaceae mericarp MC B 0 Chamerion angustifolium (L.) H o l u b Onagraceae seed PC P 2 17

12 Chelidonium majus L. Papaveraceae seed PC P 1 Chenopodium album L. Chenopodiaceae achene MC A 0 Chenopodium bonus henricus L. Chenopodiaceae achene PC P 1 Chenopodium glaucum L. Chenopodiaceae achene MC A 0 Chenopodium hybridum L. Chenopodiaceae achene MC A 0 Chenopodium pedunculare B e r t o l. Chenopodiaceae achene MC A 0 Chenopodium polyspermum L. Chenopodiaceae achene MC A 0 Chenopodium rubrum L. Chenopodiaceae achene MC A 0 Chenopodium strictum R o t h Chenopodiaceae achene MC A 0 Chenopodium suecicum M u r r Chenopodiaceae achene MC A 0 Cichorium intybus L. Asteraceae achene PC P 0 Cicuta virosa L. Apiaceae mericarp PC P 1 Circaea luteciana L. Onagraceae seed PC P 2 Cirsium arvense (L.) S c o p. Asteraceae achene PC P 2 Cirsium heterophyllum (L.) H i l l Asteraceae achene PC P 2 Cirsium oleraceum (L.) S c o p. Asteraceae achene PC P 2 Cirsium palustre (L.) S c o p. Asteraceae achene MC B 1 Cirsium vulgare (S a v i) T e n. Asteraceae achene MC B 2 Clinopodium vulgare L. Lamiaceae nutlet PC P 1 Colchicum autumnale L. Liliaceae seed PC P 0 Conium maculatum L. Apiaceae mericarp MC B 0 Consolida regalis G r a y Ranunculaceae seed MC A 0 Convallaria majalis L. Liliaceae seed PC P 2 Convolvulus arvensis L. Convolvulaceae seed PC P 2 Conyza canadensis (L.) C r o n g u i s t Asteraceae achene MC A 0 Coronilla varia L. Fabaceae seed PC P 2 Corydalis cava (L.) S c h w e i g g. et K ö r t e Fumariaceae seed PC P 0 Corydalis solida (L.) C l a i r v. Fumariaceae seed PC P 0 Crepis biennis L. Asteraceae achene MC B 2 Cruciata glabra (L.) E h r e n d. Rubiaceae mericarp PC P 1 Cuscuta europaea L. Cuscutaceae seed MC A 0 Cyanus segetum H i l l Asteraceae achene 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 PC P 2 Cynodon dactylon (L.) P e r s. Poaceae caryopsis PC P 2 Cynoglossum officinale L. Boraginaceae nutlet MC B 0 Cynosurus cristatus L. Poaceae caryopsis PC P 1 Dactylis glomerata L. Poaceae caryopsis PC P 1 Dactylorhiza fuchsii (D r u c e) S o ó Orchidaceae seed 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 PC P 0 18

13 Danthonia decumbens (L.) D C. Poaceae caryopsis PC P 1 Daucus carota L. Apiaceae mericarp MC B 0 Deschampsia cespitosa (L.) P. B e a u v. Poaceae caryopsis PC P 1 Descurainia sophia (L.) W e b b ex P r a n t l Brassicaceae seed MC A 0 Dianthus deltoides L. Caryophyllaceae seed PC P 1 Digitalis grandiflora M i l l. Scrophulariaceae seed PC P 0 Digitalis purpurea L. Scrophulariaceae seed MC B 0 Digitaria sanguinalis (L.) S c o p. Poaceae caryopsis MC A 0 Dipsacus fullonum L. Dipsacaceae achene MC B 0 Echinochloa crus galli (L.) P. B e a u v. Poaceae caryopsis MC A 0 Echium vulgare L. Boraginaceae nutlet MC B 0 Eleocharis palustris (L.) R o e m. et S c h u l t. Cyperaceae achene PC P 2 Elytrigia repens (L.) D e s v. Poaceae caryopsis PC P 2 Epilobium ciliatum R a f. Onagraceae seed PC P 2 Epilobium hirsutum L. Onagraceae seed PC P 2 Epilobium montanum L. Onagraceae seed PC P 1 Epipactis atrorubens (Hoffm.) B e s s e r Orchidaceae seed PC P 0 Epipactis helleborine (L.) C r a n t z Orchidaceae seed PC P 0 Eragrostis minor H o s t Poaceae caryopsis MC A 0 Eriophorum angustifolium H o n c k. Cyperaceae achene PC P 2 Eriophorum vaginatum L. Cyperaceae achene PC P 2 Erodium cicutarium (L.) L' H é r. Geraniaceae mericarp MC B 0 Erophila verna (L.) C h e v a l l Brassicaceae seed MC A 0 Eupatorium cannabinum L. Asteraceae achene PC P 1 Euphrasia rostkoviana H a y n e Scrophulariaceae seed MC A 0 Euphrasia stricta J. P. W o l f f Scrophulariaceae seed MC A 0 Fallopia convolvulus (L.) Á. L ö v e Polygonaceae achene MC A 0 Fallopia dumetorum (L.) H o l u b Polygonaceae achene MC A 0 Festuca arundinacea S c h r e b. Poaceae caryopsis PC P 1 Festuca gigantea (L.) V i l l. Poaceae caryopsis PC P 2 Festuca ovina L. Poaceae caryopsis PC P 1 Festuca pratensis H u d s. Poaceae caryopsis PC P 1 Festuca rubra L. Poaceae caryopsis PC P 2 Filaginella uliginosa (L.) O p i t z Asteraceae achene MC A 0 Filipendula ulmaria (L.) M a x i m. Rosaceae follicle PC P 2 Fragaria moschata (D u c h e s n e) W e s t o n Rosaceae achene PC P 1 Fragaria vesca L. Rosaceae achene PC P 2 Fumaria officinalis L. Fumariaceae seed MC A 0 Fumaria vaillantii L o i s e l. Fumariaceae seed MC A 0 19

14 Galeobdolon luteum H u d s. emend. H o l u b Lamiaceae nutlet PC P 2 Galeopsis bifida B o e n n. Lamiaceae nutlet MC A 0 Galeopsis pubescens B e s s e r Lamiaceae nutlet MC A 0 Galeopsis speciosa M i l l. Lamiaceae nutlet MC A 0 Galeopsis tetrahit L. Lamiaceae nutlet MC A 0 Galinsoga parviflora C a v. Asteraceae achene MC A 0 Galium aparine L. Rubiaceae mericarp MC A 0 Galium mollugo L. Rubiaceae mericarp PC P 1 Galium odoratum (L.) S c o p. Rubiaceae mericarp PC P 1 Galium rotundifolium L. Rubiaceae mericarp PC P 1 Galium sylvaticum L. Rubiaceae mericarp PC P 2 Galium uliginosum L. Rubiaceae mericarp PC P 1 Galium verum L. Rubiaceae mericarp PC P 1 Genista tinctoria L. Fabaceae seed PC P 1 Geranium dissectum L. Geraniaceae mericarp MC A 0 Geranium phaeum L. Geraniaceae mericarp PC P 1 Geranium pratense L. Geraniaceae mericarp PC P 1 Geranium pusillum B u r m. f. Geraniaceae mericarp MC A 0 Geranium robertianum L. Geraniaceae mericarp MC A 0 Geranium sanguineum L. Geraniaceae mericarp PC P 2 Geum urbanum L. Rosaceae achene PC P 1 Glechoma hederacea L. Lamiaceae nutlet PC P 2 Glyceria fluitans (L.) R. B r. Poaceae caryopsis PC P 2 Glyceria maxima (H a r t m.) H o l m b. Poaceae caryopsis PC P 2 Gymnadenia conopsea (L.) R. B r. Orchidaceae seed PC P 0 Hedera helix L. Araliaceae seed PC P 0 Helianthemum grandiflorum (S c o p.) D C. Cistaceae seed PC P 1 Hepatica nobilis S c h r e b. Ranunculaceae achene PC P 0 Heracleum mantegazzianum S o m m i e r et L e v i e r Apiaceae mericarp PC P 0 Heracleum sphondylium L. Apiaceae mericarp PC P 1 Herniaria glabra L. Caryophyllaceae seed MC A 0 Hieracium laevigatum W i l l d. Asteraceae achene PC P 1 Hieracium murorum L. Asteraceae achene PC P 0 Hieracium sabaudum L. Asteraceae achene PC P 2 Hieracium umbellatum L. Asteraceae achene PC P 1 Holcus lanatus L. Poaceae caryopsis PC P 1 Holcus mollis L. Poaceae caryopsis PC P 2 Homogyne alpina (L.) C a s s. Asteraceae achene PC P 1 Hordeum jubatum L. Poaceae caryopsis MC A 0 20

15 Hordeum murinum H u d s. Poaceae caryopsis MC A 0 Hyosciamus niger L. Solanaceae seed MC B 0 Hypericum perforatum L. Hypericaceae seed PC P 1 Hypochaeris radicata L. Asteraceae achene PC P 0 Impatiens glandulifera R o y l e Balsaminaceae seed MC A 0 Impatiens noli tangere L. Balsaminaceae seed MC A 0 Impatiens parviflora D C. Balsaminaceae seed MC A 0 Iris pseudacorus L. Iridaceae seed PC P 1 Iris pumila L. Iridaceae seed PC P 1 Isopyrum thalictroides L. Ranunculaceae seed PC P 1 Iva xanthiifolia N u t t. Asteraceae achene MC A 0 Jacea pratensis L a m. Asteraceae achene PC P 1 Jasione montana L. Campanulaceae seed MC B 0 Juncus articulatus L. Juncaceae seed PC P 1 Juncus bufonius L. Juncaceae seed MC A 0 Juncus conglomeratus L. Juncaceae seed PC P 1 Juncus effusus L. Juncaceae seed PC P 1 Juncus squarrosus L. Juncaceae seed PC P 1 Juncus tenuis W i l l d. Juncaceae seed PC P 1 Knautia arvensis (L.) C o u l t. Dipsacaceae achene PC P 0 Knautia maxima (O p i z) O r t m a n n Dipsacaceae achene PC P 1 Koeleria macranta (L e d e b.) S c h u l t. Poaceae caryopsis PC P 1 Koeleria pyramidata (L a m.) P. B e a u v. Poaceae caryopsis PC P 1 Lactuca serriola L. Asteraceae achene MC B 0 Lamium album L. Lamiaceae nutlet PC P 2 Lamium amplexicaule L. Lamiaceae nutlet MC A 0 Lamium maculatum L. Lamiaceae nutlet PC P 1 Lamium purpureum L. Lamiaceae nutlet MC A 0 Lapsana communis L. Asteraceae achene PC P 0 Lathraea squamaria L. Scrophulariaceae seed PC P 0 Lathyrus pratensis L. Fabaceae seed PC P 1 Lathyrus sylvestris L. Fabaceae seed PC P 1 Lathyrus vernus (L.) B e r n h. Fabaceae seed PC P 1 Lembotropis nigricans (L.) G r i s e b. Fabaceae seed PC P 1 Leontodon autumnalis L. Asteraceae achene PC P 0 Leontodon hispidus L. Asteraceae achene PC P 1 Leonurus cardiaca L. Lamiaceae nutlet PC P 2 Lepidium campestre (L.) R. B r. Brassicaceae seed MC A 0 Lepidium ruderale L. Brassicaceae seed MC B 0 21

16 Leucanthemum ircutianum D C. Asteraceae achene PC P 1 Leucojum vernum L. Amaryllaceae seed PC P 0 Libanotis pyrenaica (L.) B o u r g. Apiaceae mericarp MC B 1 Lilium martagon L. Liliaceae seed PC P 0 Linaria genistifolia (L.) M i l l. Scrophulariaceae seed PC P 1 Linaria vulgaris M i l l. Scrophulariaceae seed PC P 1 Linum usitatissimum L. Linaceae seed MC A 0 Lithospermum arvense L. Boraginaceae nutlet MC A 0 Lolium multiflorum L a m. Poaceae caryopsis MC B 1 Lolium perenne L. Poaceae caryopsis PC P 2 Lotus corniculatus L. Fabaceae seed PC P 1 Lunaria rediviva L. Brassicaceae seed PC P 2 Lupinus polyphyllus L i n d l. Fabaceae seed PC P 1 Luzula campestris (L.) D C. Juncaceae seed PC P 1 Luzula luzuloides (L a m.) D a n d y Juncaceae seed PC P 2 Luzula multiflora (E h r h.) L e j. Juncaceae seed PC P 1 Luzula pilosa (L.) W i l l d. Juncaceae seed PC P 2 Luzula sylvatica (H u d s.) G a u d i n Juncaceae seed PC P 1 Lychnis flos cuculi L. Caryophyllaceae seed PC P 1 Lycopsis arvensis L. Boraginaceae nutlet MC A 0 Lycopus europaeus (L.) Lamiaceae nutlet PC P 1 Lysimachia nummularia L. Primulaceae seed PC P 1 Lysimachia vulgaris L. Primulaceae seed PC P 1 Lythrum salicaria L. Lythraceae seed PC P 1 Maianthemum bifolium (L.) F. W. S c h m i d t Liliaceae seed PC P 0 Malva neglecta W a l l r. Malvaceae mericarp MC B 2 Matricaria discoidea D C. Asteraceae achene MC A 0 Matricaria recutital. Asteraceae achene MC A 0 Medicago lupulina L. Fabaceae seed MC B 1 Medicago sativa L. Fabaceae seed PC P 2 Melampyrum nemorosum L. Scrophulariaceae seed MC A 0 Melampyrum pratense L. Scrophulariaceae seed MC A 0 Melandrium latifolia P o i r. Caryophyllaceae seed PC P 1 Melica nutans L. Poaceae caryopsis PC P 1 Melica transsilvanica S c h u r. Poaceae caryopsis PC P 1 Melilotus albus M e d i k. Fabaceae seed MC B 0 Melilotus officinalis (L.) P a l l. Fabaceae seed MC B 1 Mentha arvensis L. Lamiaceae nutlet PC P 2 Mentha suaveolens E h r h. Lamiaceae nutlet PC P 2 22

17 Menyathes trifoliata L. Menyanthaceae seed PC P 2 Merculialis perennis L. Euphorbiaceae seed PC P 2 Microrrhinum minus (L.) F o u r r. Scrophulariaceae seed MC A 0 Milium effusum L. Poaceae caryopsis PC P 1 Mimulus guttatus D C. Scrophulariaceae seed PC P 2 Molinia caerulea (L.) M o e n c h Poaceae caryopsis PC P 1 Moneses uniflora (L.) A. G r a y Pyrolaceae seed PC P 1 Mycelis muralis (L.) D u m o r t. Asteraceae achene PC P 1 Myosotis arvensis (L.) H i l l Boraginaceae nutlet MC A 0 Myosotis laxiflora R c h b. Boraginaceae nutlet PC P 1 Myosotis nemorosa B e s s e r Boraginaceae nutlet PC P 0 Myosotis sparsiflora M i k a n ex P o h l Boraginaceae nutlet MC A 0 Myosoton aquaticum (L.) M o e n c h Caryophyllaceae seed PC P 1 Nardus stricta L. Poaceae caryopsis PC P 0 Neslia paniculata (L.) D e s v. Brassicaceae seed MC A 0 Odontites vulgaris M o e n c h Scrophulariaceae seed MC A 0 Oenanthe aquatica (L.) P o i r. Apiaceae mericarp MC B 1 Oenothera biennis L. Oenotheraceae seed MC B 1 Omalotheca sylvatica (L.) S c h. B i p. et F. W. S c h u l t z Asteraceae achene PC P 0 Omphalodes scorpioides (H a e n k e) S c h r a n k Boraginaceae nutlet MC B 0 Onobrychis viciifolia S c o p. Fabaceae seed PC P 0 Onopordum acanthium L. Asteraceae achene MC B 1 Origanum vulgare L. Lamiaceae nutlet PC P 1 Orthantha lutea (L.) A. K e r n. ex W e t t s t. Scrophulariaceae seed MC A 0 Oxalis acetosella L. Oxalidaceae seed PC P 1 Oxalis stricta L. Oxalidaceae seed MC A 0 Oxyccocus palustris P e r s. Ericaceae seed PC P 2 Papaver dubium L. Papaveraceae seed MC A 0 Papaver rhoeas L. Papaveraceae seed MC A 0 Pastinaca sativa L. Apiaceae mericarp MC B 0 Pedicularis sylvatica L. Scrophulariaceae seed PC P 1 Pedicularis verticillata L. Scrophulariaceae seed PC P 1 Persicaria hydropiper (L.) S p a c h Polygonaceae achene MC A 1 Persicaria maculata (Rafin.) G r a y Polygonaceae achene 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 PC P 2 Peucedanum palustre (L.) M o e n c h Apiaceae mericarp PC P 1 Phalaroides arundinacea (L.) R a u s c h e r t Poaceae caryopsis PC P 2 Phleum pratense L. Poaceae caryopsis PC P 1 Phragmites australis (C a v.) T r i n. Poaceae caryopsis PC P 2 23

18 Phyteuma nigrum S c h m. Campanulaceae seed PC P 1 Pimpinella major (L.) H u d s. Apiaceae mericarp PC P 1 Pimpinella saxifraga L. Apiaceae mericarp PC P 1 Plantago lanceolata L. Plantaginaceae seed PC P 0 Plantago major L. Plantaginaceae seed PC P 0 Poa annua L. Poaceae caryopsis MC A 0 Poa compressa L. Poaceae caryopsis PC P 1 Poa nemoralis L. Poaceae caryopsis PC P 2 Poa pratensis L. Poaceae caryopsis PC P 1 Polygonatum odoratum (M i l l.) D r u c e Liliaceae seed PC P 1 Polygonum aviculare L. Polygonaceae achene MC A 0 Polygonum lapathifolia (L.) G r a y Polygonaceae achene MC A 0 Portulaca oleracea L. Portulacaceae seed MC A 0 Potamogeton pectinatus L. Potamogetonaceae achene PC P 2 Potentilla argentea L. Rosaceae achene PC P 1 Potentilla erecta (L.) R d e u s c h. Rosaceae achene PC P 2 Potentilla recta L. Rosaceae achene PC P 0 Potentilla reptans L. Rosaceae achene PC P 1 Potentilla supina L. Rosaceae achene MC B 1 Potentilla tabernaemontani A s c h. Rosaceae achene PC P 0 Prenanthes purpurea L. Asteraceae achene PC P 2 Primula elatior (L.) H i l l. Primulaceae seed PC P 0 Primula veris L. Primulaceae seed PC P 1 Prunella vulgaris L. Lamiaceae nutlet PC P 1 Pseudofumaria lutea (L.) M e d i k. Fumariaceae seed PC P 0 Pseudolysimachion spicatum (L.) O p i z Scrophulariaceae seed PC P 0 Puccinellia distans (J a c q.) P a r l. Poaceae caryopsis PC P 0 Pulmonaria officinalis L. Boraginaceae nutlet PC P 0 Pulsatilla grandis W e n d e r. Ranunculaceae achene PC P 0 Pyrethrum corymbosum (L.) S c o p. Asteraceae achene PC P 0 Pyrola minor L. Pyrolaceae seed PC P 1 Ranunculus acris L. Ranunculaceae achene PC P 0 Ranunculus auricomus L. Ranunculaceae achene PC P 0 Ranunculus flammula L. Ranunculaceae achene PC P 2 Ranunculus sceleratus L. Ranunculaceae achene MC A 0 Raphanus raphanistrum L. Brassicaceae seed MC A 0 Reseda lutea L. Resedaceae seed MC B 1 Reseda luteola L. Resedaceae seed MC B 1 Rhinanthus minor L. Scrophulariaceae seed MC A 0 24

19 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 MC A 0 Rhinanthus serotinus (S c h ö n h.) O b o r n y Scrophulariaceae seed MC A 0 Rorippa palustris (L.) B e s s e r Brassicaceae seed MC B 1 Rorippa sylvestris (L.) B e s s e r Brassicaceae seed PC P 1 Rumex crispus L. Polygonaceae achene PC P 0 Rumex maritimus L. Polygonaceae achene MC A 0 Rumex obtusifolius L. Polygonaceae achene PC P 0 Sagina procumbens L. Caryophyllaceae seed PC P 0 Salvia pratensis L. Lamiaceae nutlet PC P 1 Sanguisorba minor S c o p. Rosaceae achene PC P 0 Sanguisorba officinalis L. Rosaceae achene PC P 1 Sanicula europaea L. Apiaceae mericarp PC P 1 Saponaria officinalis L. Caryophyllaceae seed PC P 1 Saxifraga granulata L. Saxifragaceae seed PC P 0 Scirpus sylvaticus L. Cyperaceae achene PC P 2 Scorzonera humilis L. Asteraceae achene PC P 0 Scrophularia nodosa L. Scrophulariaceae seed PC P 0 Scutellaria galericulata L. Lamiaceae nutlet PC P 2 Sedum acre L. Crassulaceae seed PC P 0 Selinum carvifolia (L.) L. Apiaceae mericarp PC P 0 Senecio erraticus B e r t o l. Asteraceae achene 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 PC P 2 Senecio rivularis (W. K.) D C. Asteraceae achene PC P 2 Senecio sylvaticus L. Asteraceae achene MC A 0 Senecio viscosus L. Asteraceae achene MC A 0 Senecio vulgaris L. Asteraceae achene MC A 0 Setaria pumila (P o i r.) R o e m. et S c h u l t. Poaceae caryopsis MC A 0 Silaum silaus (L.) S c h i n z et T h e l l. Apiaceae mericarp PC P 1 Silene dioica (L.) C l a i r v. Caryophyllaceae seed PC P 1 Silene nutans L. Caryophyllaceae seed PC P 1 Sinapis alba L. Brassicaceae seed MC A 0 Sinapis arvensis L. Brassicaceae seed MC A 0 Sisymbrium altissimum L. Brassicaceae seed MC A 0 Sisymbrium loeselii L. Brassicaceae seed MC A 0 Sisymbrium officinale (L.) S c o p. Brassicaceae seed MC A 0 Solanum dulcamara L. Solanaceae seed PC P 2 Solanum nigrum L. Solanaceae seed MC A 0 Solanum vilosum (L.) M i l l. Solanaceae seed MC A 0 Soldanella montana W i l l d. Primulaceae seed PC P 0 25

20 Solidago gigantea A i t o n Asteraceae achene PC P 2 Solidago virgaurea L. Asteraceae achene PC P 1 Sonchus arvensis L. Asteraceae achene PC P 1 Sonchus oleraceus L. Asteraceae achene MC A 0 Sparganium erectum L. Sparganiaceae seed PC P 2 Spergula arvensis L. Caryophyllaceae seed MC A 0 Spergularia rubra (L.) J. P r e s l et C. P r e s l Caryophyllaceae seed MC A 0 Stachys palustris L. Lamiaceae nutlet PC P 2 Stachys sylvatica L. Lamiaceae nutlet PC P 1 Stellaria graminea L. Caryophyllaceae seed PC P 1 Stellaria holostea L. Caryophyllaceae seed PC P 1 Steris viscaria (L.) R a f i n. Caryophyllaceae seed PC P 1 Symphytum tuberosum L. Boraginaceae nutlet PC P 1 Tanacetum vulgare L. Asteraceae achene PC P 2 Taraxacum sect. Ruderalia K i r s c h n e r et al. Asteraceae achene PC P 0 Teucrium chamaedrys L. Lamiaceae nutlet PC P 1 Thalictrum lucidum L. Ranunculaceae achene PC P 1 Thlaspi arvense L. Brassicaceae seed MC A 0 Thlaspi perfoliatum L. Brassicaceae seed MC A 0 Thymus pulegioides L. Lamiaceae nutlet PC P 1 Tofieldia calyculata (L.) W a h l e n b. Liliaceae seed PC P 1 Torilis japonica (Houtt.) D C. Apiaceae mericarp MC B 0 Tragopogon pratensis L. Asteraceae achene PC P 1 Trientalis europaea L. Primulaceae seed PC P 0 Trifolium alpestre L. Fabaceae seed PC P 1 Trifolium arvense L. Fabaceae seed MC A 0 Trifolium aureum P o l l i c h Fabaceae seed MC A 0 Trifolium campestre S c h r e b. Fabaceae seed MC A 0 Trifolium dubium S i b t h. Fabaceae seed MC A 0 Trifolium medium G r u f b. Fabaceae seed PC P 1 Trifolium repens L. Fabaceae seed PC P 2 Trifolium spadiceum L. Fabaceae seed MC A 0 Tripleurospermum perforatum (M é r a t) M. L a í n z Asteraceae achene MC A 0 Trisetum flavescens (L.) P. B e a u v. Poaceae caryopsis PC P 1 Tussilago farfara L. Asteraceae achene PC P 2 Typha latifolia L. Typhaceae follicle PC P 2 Tythymalus cyparissias (L.) S c o p. Euphorbiaceae seed PC P 2 Tythymalus esula (L.) H i l l. Euphorbiaceae seed PC P 1 Tythymalus helioscopia (L.) H i l l. Euphorbiaceae seed MC A 0 26

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