Plant heights are relevant for the Czech Republic. They are measured in metres and relate to fully developed mature generative plants growing in the wild. Each taxon is characterized by two values: minimum (lower limit of the common range) and maximum (upper limit of the common range). The data were taken from the Key to the Flora of the Czech Republic (Kaplan et al. 2019).
Kaplan Z., Danihelka J., Chrtek J. Jr., Kirschner J., Kubát K., Štěpánek J. & Štech M. (eds) (2019) Klíč ke květeně České republiky [Key to the flora of the Czech Republic]. Ed. 2. – Academia, Praha.
Growth form describes the potential life span of the plant and its parts (ramets), its reproductive strategy and durability of its aboveground parts (Klimešová et al. 2016, Ottaviani et al. 2017). Here the growth form is classified into nine categories, which also consider herbaceous vs woody nature of the stem. Annual herbs live for one season only and reproduce by seed usually in the same season in which they germinated. They may but need not be clonal; their clonality typically does not result in fragmentation. Perennial herbs are divided into three categories: (i) monocarpic perennial non-clonal herbs, which reproduce sexually only once in their life and do not possess woody aboveground parts or organs of clonal growth, (ii) polycarpic perennial non-clonal herbs, which reproduce sexually several times during their life and do not possess organs of clonal growth, and (iii) clonal herbs, which possess organs of clonal growth enabling them to make fragments during their life and to form independent units (ramets) by vegetative reproduction; the whole plant reproduces sexually several times during its life, while individual ramets may reproduce once or several times during their life. The other categories include woody plants, which may but need not possess organs of clonal growth and may be able or not of fragmentation and vegetative reproduction. The woody plants are divided into dwarf shrubs (woody plants lower than 30 cm, also including suffruticose plants with erect, herbaceous shoots growing from woody stems at the base, which die out in autumn except for the lowest part with regenerative buds), shrubs (woody plants higher than 30 cm, branched at the base), trees (woody plants with trunk and crown), woody lianas and parasitic epiphytes, which include only two species of the Czech flora, Loranthus europaeus and Viscum album.
Data were partly taken from the aggregated CLO-PLA 3.4 database (Klimešová et al. 2017). The CLO-PLA categories were further divided into separate categories for herbaceous vs woody plants, and taxa not included in CLO-PLA were added.
Dřevojan P. (2020) Growth form. – www.pladias.cz.
Klimešová J., Nobis M. P. & Herben T. (2016) Links between shoot and plant longevity and plant economics
spectrum: Environmental and demographic implications. – Perspectives in Plant Ecology, Evolution and
Systematics 22: 55–62.
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Ottaviani G., Martínková J., Herben T., Pausas J. G. & Klimešová J. (2017) On plant modularity traits: functions
and challenges. – Trends in Plant Science 22: 648–651.
Life form classification follows the system of Raunkiaer (1934), which is based on the position of the buds that survive the unfavourable season. Macrophanerophytes are woody plants that bear the surviving buds at least 2 m above the ground, usually trees; nanophanerophytes are woody plants with surviving buds 0.3–2 m above the ground, usually shrubs; chamaephytes are herbs or low woody plants with surviving buds above the ground, but not more than 30 cm above it; hemicryptophytes are perennial or biennial herbs with surviving buds on aboveground shoots at the level of the ground; geophytes are perennial plants with surviving buds belowground, usually with bulbs, tubers or rhizomes; hydrophytes are plants with surviving buds in water, usually on the bottom of water bodies; therophytes are summer- or winter-annual herbs that survive the unfavourable season only as seeds germinating in autumn, winter or spring.
The data on life forms were taken from the Key to the Flora of the Czech Republic (Kaplan et al. 2019). Newly added alien taxa were assigned to the categories of life forms based on the FloraVeg.EU database (Dřevojan et al. 2022). Some taxa can belong to more than one life form. In such cases, the dominant life form is listed first.
Kaplan Z., Danihelka J., Chrtek J. Jr., Kirschner J., Kubát K., Štěpánek J. & Štech M. (eds) (2019) Klíč ke květeně České republiky [Key to the flora of the Czech Republic]. Ed. 2. – Academia, Praha.
Dřevojan P., Čeplová N., Štěpánková P. & Axmanová I. (2022) Life form. – www.FloraVeg.EU.
Raunkiaer C. (1934) The life forms of plants and statistical plant geography. – Clarendon Press, Oxford.
Grime (1974, 1979) distinguished three basic ecological strategies of plants: competitive strategy (C), advantageous in habitats where resources are abundant, conditions not extreme and disturbance level is low; stress tolerant strategy (S), advantageous where resources are scarce, conditions severe, but disturbance is uncommon; and ruderal strategy (R), advantageous where resources are abundant and conditions not extreme, but disturbance level is high. There are also intermediate strategies in all possible combinations of the three basic types (CR, CS, SR, CSR). Data were taken from the BiolFlor database (Klotz & Kühn 2002).
Klotz S. & Kühn I. (2002) Ökologische Strategietypen. – In: Klotz S., Kühn I. & Durka W. (eds), BIOLFLOR: eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland, Schriftenreihe für Vegetationskunde 38: 119–126.
Grime J. P. (1974) Vegetation classification by reference to strategies. – Nature 250: 26–31.
Grime J. P. (1979) Plant strategies and vegetation processes. – Wiley, Chichester.
Data on the presence of leaves on the plant, their metamorphoses and reductions are based on the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010) and the Key to the Flora of the Czech Republic (Kubát et al. 2002).
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky
[Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Kubát K., Hrouda L., Chrtek J. Jr., Kaplan Z., Kirschner J. & Štěpánek J. (eds) (2002) Klíč ke květeně České
republiky [Key to the flora of the Czech Republic]. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Four basic types of leaf arrangement are distinguished: alternate, opposite, verticillate (whorled) and rosulate (in the basal rosette). The character is assessed in well-developed plants, i.e. not in individuals re-sprouting after damage by mowing or grazing or those with teratological modifications. More than one character state may occur (e.g. Hylotelephium jullianum and Salix purpurea) in some taxa: all character states are recorded in such cases.
In some plants, the arrangement of frondose bracts in the inflorescence is assessed separately (e.g. true leaves in Veronica persica and V. polita are opposite, while bracts are alternate). Leaves with interpetiolar stipules found in the Rubiaceae family are considered as whorled. The leaves in Rhamnus cathartica are considered as opposite, although in most cases they are sub-opposite.
The information was extracted mainly from the descriptions in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). In cases of uncertainties, mainly for alien taxa, additional sources were consulted, including the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (www.tropicos.org/Project/Pakistan).
Grulich V., Holubová D., Štěpánková P. & Řezníčková M. (2017) Leaf arrangement. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
The primary distinction is made between simple and compound leaves. The simple leaves are categorized based on the leaf blade division associated with venation into palmately divided (e.g. Alchemilla), pinnately divided (e.g. Achillea millefolium), forked (e.g. Batrachium, Ceratophyllum and Utricularia) and pedate (e.g. Helleborus). The categorization is based on well-developed leaves. In many taxa, transitions occur between simple leaves with a dentate or serrate margin, and simple divided (pinnately or palmately lobed) leaves. Only the leaves with the lamina divided to at least one-quarter of their width are considered as divided. Many taxa with varying leaf division are assigned to more than one character state.
The compound leaves are divided into palmate and pinnate. The taxa that have both ternate and pinnate leaves, the latter with two pairs of leaflets (e.g. Aegopodium podagraria and some other species of the Apiaceae family), are assigned to both character states. The degree of division in pinnately compound leaves indicated here relates to well-developed leaves, especially to the basal part of the lamina. Taxa with multiple pinnately compound leaves are assigned to two or more character states based on the level of division, but very small leaves, which may correspond to simple leaves, are not considered.
In many cases, there are transitions between simple and compound leaves, especially between pinnatisect and pinnate leaves. Leaves with linear or filiform segments, including the bi-, tri- or even more-pinnatisect or palmatisect leaves (e.g. stem leaves in Batrachium fluitans, Cardamine pratensis and the genus Seseli) are classified as simple (dissected) leaves. In contrast, leaves with broader segments attached to the rachis by a distinct constriction or a petiolule (e.g. stem leaves in Cardamine dentata or ground leaves in Pimpinella saxifraga) are classified as compound.
In heterophyllous taxa, all types of leaves are assessed, and the taxon is assigned to two or more character states. However, less divided leaves found in juvenile plants of some taxa are not considered heterophyllous. The parasitic plants with rudimentary (vestigial) leaves (e.g. Cuscuta) or the plants with phylloclades replacing the vestigial leaves (e.g. Asparagus) are assigned the character state “reduced”.
The information was extracted mainly from the descriptions in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). In uncertain cases, mainly for alien taxa, additional sources were consulted, including the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (www.tropicos.org/Project/Pakistan).
Grulich V., Holubová D., Štěpánková P. & Řezníčková M. (2017) Leaf shape. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
Stipules, i.e. paired leaflike appendages at the base of the petiole or sessile leaf blade, can be present or absent. Caducous stipules, i.e. those disappearing soon after the leaf blade has developed (e.g. Prunus), are considered as present. The interpetiolar stipules, morphologically indistinguishable from true leaves and together forming whorls (e.g. Rubiaceae), are considered as true stipules. In contrast, stipules modified into glands (e.g. Lotus) or hairs (e.g. Portulacaceae) are not considered as stipules here.
Information about the presence of stipules was extracted from the descriptions in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). In cases of uncertainties, mainly concerning alien taxa, descriptions in the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (www.tropicos.org/Project/Pakistan) were consulted.
Grulich V., Holubová D., Štěpánková P. & Řezníčková M. (2017) Stipules. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
Leaf petiole can be present or absent. In some plants, it can be present in some leaves but absent in others. The data were extracted from the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010), the Key to the Flora of the Czech Republic (Kubát et al. 2002), the New Hungarian Herbal (Király et al. 2011) and the Excursion Flora of Germany (Jäger & Werner 2000).
Prokešová H. & Grulich V. (2017) Petiole. – www.pladias.cz.
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Jäger E. J. & Werner K. (eds) (2000) Rothmaler, Exkursionsflora von Deutschland. Band 3. Gefäßpflanzen: Atlasband. Ed. 10. – Spectrum Akademischer Verlag, Heidelberg & Berlin.
Király G., Virók V. & Molnár V. (eds) (2011) Új Magyar füvészkönyv. Magyarország hajtásos növényei: ábrák [New Hungarian Herbal. The vascular plants of Hungary: Figures]. – Aggteleki Nemzeti Park Igazgatóság, Jósvafő.
Kubát K., Hrouda L., Chrtek J. Jr., Kaplan Z., Kirschner J. & Štěpánek J. (eds) (2002) Klíč ke květeně České republiky [Key to the flora of the Czech Republic]. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Leaf life span is a functional trait important for plant competitiveness. It depends on the climate in the distribution range of the taxon and microclimate, nutrient and light availability in typical habitats of the taxon. The data were taken from the BiolFlor database (Klotz & Kühn 2002).
Categories
Klotz S. & Kühn I. (2002) Blattmerkmale. – In: Klotz S., Kühn I. & Durka W. (eds), BIOLFLOR: eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland, Schriftenreihe für Vegetationskunde 38: 119–126.
The months of the beginning and end of flowering in the Czech Republic are given. The data were taken from the Key to the Flora of the Czech Republic (Kaplan et al. 2019).
Kaplan Z., Danihelka J., Chrtek J. Jr., Kirschner J., Kubát K., Štěpánek J. & Štech M. (eds) (2019) Klíč ke květeně České republiky [Key to the flora of the Czech Republic]. Ed. 2. – Academia, Praha.
Flower colour is reported for nearly all angiosperms except duckweeds (Araceae p. p.) and some hybrids for which data on flower colour were not available.
If a species has more than one flower colour, all colours are reported irrespective of their frequency. This approach is used both for species that regularly form populations with different flower colours (e.g. Corydalis cava and Iris pumila) and for species with occasional occurrence of deviating flower colour (e.g. albinism in Salvia pratensis or pink flowers in Ajuga reptans). However, the whole range of variation is not fully reported in cultivated plants, for which some cultivars of different colour may be ignored (e.g. Gladiolus hortulanus and Callistephus chinensis). In plants with flowers of two colours (e.g. Cypripedium calceolus), both colours are reported. In plants with multi-coloured flowers (e.g. the variegated lip in Ophrys apifera) the predominant colour is reported.
If the flower has a well-developed perianth, the reported flower colour relates to the corolla or the tepals of the homochlamydeous perianth. If such a flower has bracts of a contrasting colour (e.g. Melampyrum nemorosum), their colour is not considered. If the corolla or the homochlamydeous perianth is not developed, the flower colour is based on the calyx (e.g. Daphne mezereum), bracts (e.g. Aristolochia clematitis), the system of bracts and bracteoles in the inflorescence (Euphorbia) or the involucre on secondary peduncles (Bupleurum longifolium). In species of Araceae with spadix and spathe of contrasting colours (e.g. Calla palustris) both colours are reported. The colour of the whole inflorescence is reported for some plants with reduced flowers (e.g. Betula, Salix, some Cyperaceae and Typhaceae). Spikelets in Poaceae are reported as green disregarding a possible violet tint; exceptions include the Melica ciliata agg. and Cortaderia that are reported as white. Also in other, rare cases, the inflorescence colour is reported as flower colour (e.g. green in Ficus carica). In Asteraceae, the colours of the disk flowers and ray flowers are reported separately if the ray flowers are developed and have a contrasting colour (e.g. Bellis perennis). The colour of the involucrum is reported for species with tiny flower heads and indistinct flowers (e.g. Artemisia campestris and Xanthium) and for “immortelles” (e.g. Helichrysum and Xeranthemum).
Information on flower colour is partly based on the field knowledge, partly obtained from various photographs and descriptions in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). In the taxa that are not reported in the Flora of the Czech Republic, as well as in unclear cases (especially in alien species), other sources were used, especially the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (http://www.tropicos.org/Project/Pakistan).
Categories
Štěpánková P. & Grulich V. (2019) Flower colour. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
Flowers of angiosperms are either zygomorphic (with bilateral symmetry) or actinomorphic (with radial symmetry). This character is not reported for taxa with achlamydeous flowers and taxa with strongly reduced or rudimentary perianth or with a perianth modified into scale-like or setaceous structures. However, it is reported for taxa with the perianth reduced to a corolla-like calyx (e.g. Aizoaceae and Daphne) and in taxa with flowers surrounded by complex structures combining bracts with the proper perianth or petal-like staminodes and stamens (e.g. Canna). Spiral and spirocyclic flowers, though actually asymmetric, are classified as actinomorphic in Nymphaeaceae and most species of Ranunculaceae. In contrast, in some other members of Ranunculaceae (e.g. Aconitum and Delphinium), they are classified as zygomorphic. Bisymmetric flowers (in the Brassicaceae family and the genera Dicentra and Lamprocapnos) are consistently classified as actinomorphic. Both zygomorphic and actinomorphic flowers are reported for taxa with both symmetry types (e.g. Succisa pratensis).
The information about flower symmetry was extracted from the descriptions in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). If some uncertainty occurred, particularly in some alien taxa, the descriptions in the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (www.tropicos.org/Project/Pakistan) were consulted.
Grulich V., Holubová D., Štěpánková P. & Řezníčková M. (2017) Flower symmetry. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
Perianth (perigon), i.e. the non-reproductive part of the angiosperm flower, can be classified into heterochlamydeous and homochlamydeous. Heterochlamydeous flowers are divided into calyx and corolla. In homochlamydeous flowers, calyx and corolla are indistinguishable. Perianth or some of its parts can be reduced or absent; flowers with no perianth are called achlamydeous.
In Apiaceae, the presence of the calyx teeth is assessed as a reduced calyx; if these teeth are not visible, the calyx is considered as absent. In Asteraceae, the presence of a pappus, scales or a collar-like structure is considered as a reduced calyx; if no such structures are present, the calyx is considered as absent. In Cyperaceae, the presence of perianth bristles is assessed as a reduced perianth. All members of the Poaceae family are considered as plants with a reduced perianth. The perianth in the genus Basella is arbitrarily classified as a reduced calyx though it is also often considered as a reduced homochlamydeous perianth. The character states “homochlamydeous, sometimes absent” and “homochlamydeous, reduced or absent” mean that in one plant some flowers may have a well-developed or reduced perianth, while other flowers may be achlamydeous (e.g. Atriplex).
The information was extracted mainly from the descriptions in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). For the taxa not treated in that flora or if uncertainties occurred, mainly concerning some alien taxa, the descriptions in the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (www.tropicos.org/Project/Pakistan) were consulted.
Grulich V., Prokešová H. & Štěpánková P. (2017) Perianth type. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
This characteristic of angiosperm flowers is assessed either as a fusion of the corolla or, in homochlamydeous taxa (e.g. Amaryllidaceae, Liliaceae and Orchidaceae) as a fusion of the whole perianth. It is not assessed in achlamydeous groups (e.g. Salix) and plants with a strongly reduced or rudimentary perianth or with the perianth modified in scale-like or setaceous structures with a varying number of bristles, which may be free (e.g. in Cyperaceae) or partially fused (e.g. in most of Poaceae). The perianth of such plants is considered as reduced. The perianth in the genus Aristolochia is also classified as reduced (neither fused nor free): it is modified to scales situated at the bottom of a tube-like structure formed by fused bracts. Both primary character states are assigned to the taxa with unisexual male and female flowers that differ in the fusion of the perianth (e.g. Cannabis). A similar approach is used in the taxa in which some flowers are homochlamydeous while others are achlamydeous (e.g. Atriplex).
The basic information was extracted from the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). If some uncertainty occurred, especially for alien taxa, other sources were consulted, including the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (www.tropicos.org/Project/Pakistan).
Grulich V., Holubová D., Štěpánková P. & Řezníčková M. (2017) Perianth fusion. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
This characteristic of angiosperm flowers is not assessed for achlamydeous groups (e.g. Salix) and plants with a strongly reduced or rudimentary perianth or with the perianth modified in scale-like or setaceous structures (e.g. Cyperaceae and Poaceae). In Amaranthaceae and the genus Cannabis, the perianth is recognizable, and the degree of its fusion could be assessed, but not its shape. If the corolla or the perianth have an intermediate shape between two character states, the taxon is assigned to both of them. Many sympetalous corollas and syntepalous perianths have unique shapes that are difficult to match to general classification categories. The taxa with such shapes are classified to an auxiliary category “special type” (e.g. Canna, Cyclamen, Dicentra, Gladiolus, Impatiens and Iris).
The basic information was extracted from the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). In uncertain cases, especially in some alien taxa, other sources were consulted, including the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (www.tropicos.org/Project/Pakistan).
Grulich V., Holubová D., Štěpánková P. & Řezníčková M. (2017) Shape of the sympetalous corolla or syntepalous perianth. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
The calyx of angiosperm flowers can be fused into a calyx tube (synsepalous calyx) or composed of distinct sepals (aposepalous). In some plants (especially in Asteraceae) the calyx is modified into a ring of fine feathery hairs called the pappus. Taxa with both synsepalous and aposepalous calyx (e.g. Platanus) are classified as “synsepalous and aposepalous”. A cup-shaped tube formed of fused sepals, petals and stamens is called hypanthium. However, hypanthium may also be interpreted as a product of an intercalary growth of the floral axis (receptacle) up and around the carpels, forming a cup-shaped structure, sometimes even fusing with the outer walls of the carpels and making the ovary inferior. In most genera of the Onagraceae family, the hypanthium forms a floral tube fairly overtopping the apex of the ovary.
The data were taken from the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010), the Key to the Flora of the Czech Republic (Kubát et al. 2002), the New Hungarian Herbal (Király et al. 2011) and the Excursion Flora of Germany (Jäger & Werner 2000).
Prokešová H. & Grulich V. (2017) Calyx fusion. – www.pladias.cz.
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Jäger E. J. & Werner K. (eds) (2000) Rothmaler, Exkursionsflora von Deutschland. Band 3. Gefäßpflanzen: Atlasband. Ed. 10. – Spectrum Akademischer Verlag, Heidelberg & Berlin.
Király G., Virók V. & Molnár V. (eds) (2011) Új Magyar füvészkönyv. Magyarország hajtásos növényei: ábrák [New Hungarian Herbal. The vascular plants of Hungary: Figures]. – Aggteleki Nemzeti Park Igazgatóság, Jósvafő.
Kubát K., Hrouda L., Chrtek J. Jr., Kaplan Z., Kirschner J. & Štěpánek J. (eds) (2002) Klíč ke květeně České republiky [Key to the flora of the Czech Republic]. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Inflorescence types follow the morphological system used in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). As the Czech terminology used for inflorescences does not match the English terminology, we use Latin terms in the English version of the Pladias Database. The exact identification of the inflorescence type is often equivocal because of varying interpretations of the same object. In species with unisexual flowers, male and female flowers can occur in different inflorescence types. In other cases, it is not possible to identify the inflorescence without detailed knowledge of evolutionary morphology, e.g. umbella vs pseudumbella in the genus Butomus. There are also compound inflorescences, in some cases with very different structure of their parts, especially in Asteraceae, which can have even triple inflorescences (e.g. Echinops sphaerocephalus often has an anthella ex capitulis anthodiorum composita).
The information was extracted mainly from the descriptions in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). For the taxa not treated in that flora or if some uncertainties occurred, mainly concerning some alien taxa, information was taken from the descriptions in the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994) and the Flora of Pakistan (www.tropicos.org/Project/Pakistan). In critical groups (e.g. Rubus), especially in recently described species, inflorescence type was taken from the original descriptions.
Grulich V. & Štěpánková P. (2019) Inflorescence type. – www.pladias.cz.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical
Garden, St. Louis.
Dicliny characterizes the level of spatial separation of male and female reproductive organs. Monoclinous (synoecious) plants, including most taxa of the central-European flora, have only bisexual (hermaphroditic) flowers. The plants with unisexual flowers are either monoecious (with both male and female flowers growing on the same individual) or dioecious (with male and female flowers growing on different individuals). Gynomonoecious plants have female and bisexual flowers on the same individuals, while andromonoecious plants have male and bisexual flowers on the same individuals. Gynodioecious plants have female and bisexual flowers on different individuals, or some individuals have only female flowers, and other individuals have both male and female flowers. Androdioecious plants have male and bisexual flowers on different individuals, or some individuals have only male flowers, and other individuals have both male and female flowers. Trioecious plants have individuals with male flowers, individuals with female flowers, and individuals with bisexual (or both male and female unisexual) flowers. Trimonoecious plants have a male, female and bisexual flowers on the same individual. Other plants can be male sterile. The data on dicliny were taken from the BiolFlor database (Durka 2002).
Durka W. (2002) Blüten- und Reproduktionsbiologie. – In: Klotz S., Kühn I. & Durka W. (eds), BIOLFLOR – Eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland, Schriftenreihe für Vegetationskunde 38: 133–175.
The type of generative reproduction (breeding system) is defined by the origin of gametes that fuse to form offspring. On the one hand, it includes obligate outcrossing, which can be controlled by genetic mechanisms of recognition and rejection of self-pollen before the fertilization of the egg cell (allogamy, self-incompatibility), sequential hermaphroditism (dichogamy) or unisexuality of plant individuals (dioecy). On the other hand, it includes obligate autogamy, which refers to the fusion of two gametes that both originate from one flower or one individual. However, various mixed strategies are common, including reproduction by both self-fertilization and mating with other individuals. The degree of self-fertilization can be affected by both genetic and ecological factors, among others by frequency, diversity and foraging strategy of pollinators. Three categories are distinguished: (i) facultative allogamy (outcrossing prevails, but selfing is possible), (ii) facultative autogamy (mainly selfing, outcrossing is rare) and (iii) mixed mating, in which both outcrossing and selfing are common, sometimes with different frequencies among populations. The last main category, apomixis, includes seed production without fertilization. It can be either obligate (offspring is genetically identical with the maternal plant) or facultative (accompanied by residual sexuality, as a rule with a low frequency). Hybrid plants are often sterile, but can sometimes reproduce vegetatively and persist for a long time. In some cases (e.g. Pilosella), such sterile hybrids are considered as species and included in this list. Some morphologically well-defined and widely accepted taxa consist of populations with contrasting modes of reproduction (as a rule connected with ploidy levels). For example, some populations are sexual and allogamous while others are apomictic. Data on generative reproduction types of the taxa of Czech flora were obtained through a search of the available literature.
Chrtek J. Jr. (2018) Generative reproduction type. – www.pladias.cz.
Pollen is transferred to stigma by different vectors, including abiotic vectors such as wind (anemophily) or water (hydrophily), or biotic vectors such as insects (entomophily). An alternative mechanism is selfing (autogamy), which can include special mechanisms such as cleistogamy (selfing in rudimentary, obligatorily autogamous flowers), pseudocleistogamy (selfing in flowers that do not open due to adverse environmental conditions) or geitonogamy (selfing by pollen from a neighbouring flower of the same plant except the cases of pollen transfer by a vector). Pollination syndromes are adopted from the BiolFlor database (Durka 2002).
Durka W. (2002) Blüten- und Reproduktionsbiologie. – In: Klotz S., Kühn I. & Durka W. (eds), BIOLFLOR – Eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland, Schriftenreihe für Vegetationskunde 38: 133–175.
The primary classification of fruit types is into dry and fleshy. Within each of these two groups, fruit types are further classified based on the scheme outlined in the first volume of the Flora of the Czech Republic (Slavíková 1988), which consistently uses the typological method. This means that fruits are classified based purely on their morphology following the formal definitions of the fruit type, regardless of the fruit type found in closely related species or genera.
One-seeded fruits in Brassicaceae (e.g. Crambe) are classified as achenes, not siliculas. Indehiscent two- and more-seeded fruits in the same family, breaking mainly in constrictions (e.g. in Bunias and Raphanus), are consistently classified as a loment, even if the fruit breaks into two distinct parts, of which one is one-seeded and the other, of strikingly different shape, two- or more-seeded and dehiscent, such as in Rapistrum rugosum. A similar approach is used for the classification of fruits in Fabaceae. Dehiscent fruits of most taxa are classified as legumes, while indehiscent two- and more-seeded fruits breaking into single-seeded parts (e.g. in Hippocrepis and Securigera) are classified as loments. One-seeded indehiscent fruits (e.g. in Onobrychis and Trifolium) are classified as achenes. Two- or more-seeded indehiscent fruits (e.g. in Sophora japonica and Vicia faba) are also classified as legumes. The fruits of all Euphorbia species are classified as capsules, although in some cases the seeds are not released. Fleshy false fruits of the genera Basella, Ficus, Maclura, Morus, Nuphar and Nymphaea are merged into a separate category.
The information about fruit type was extracted mainly from the descriptions in the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010). For the taxa not treated in that flora or in case of uncertainties, especially regarding alien taxa, descriptions in the Flora of North America (Flora of North America Editorial Committee 1993), the Flora of China (Wu et al. 1994), the Flora of Pakistan (www.tropicos.org/Project/Pakistan), and Flora Iberica (Castroviejo et al. 1986; mainly for the Fabaceae family) were consulted.
Grulich V., Holubová D., Štěpánková P. & Řezníčková M. (2017) Fruit type. – www.pladias.cz.
Castroviejo S., Laínz M., López González G., Montserrat P., Muńoz Garmendia F., Paiva J. & Villar L. (eds) (1986) Flora Iberica. Plantas vasculares de la Península Ibérica e Islas Baleares. – Real Jardín Botánico, Madrid.
Flora of North America Editorial Committee (eds) (1993) Flora of North America North of Mexico. – Oxford
University Press, New York.
Flora of Pakistan. – http://www.tropicos.org/Project/Pakistan
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Slavíková Z. (1988) Terminologický slovník [Terminological dictionary]. – In: Hejný S., Slavík B., Chrtek J.,
Tomšovic P. & Kovanda M. (eds), Květena České socialistické republiky [Flora of the Czech Socialist
Republic] 1: 130–153, Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Wu Z., Raven P. H. & Huang D. (eds) (1994) Flora of China. – Science Press, Beijing & Missouri Botanical Garden, St. Louis.
Data on fruit colour according to the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010) and the Key to the Flora of the Czech Republic (Kubát et al. 2002). Fruit colours are standardized into ten colours. A single dominant colour of ripe fruit is reported for each taxon.
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky
[Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Kubát K., Hrouda L., Chrtek J. Jr., Kaplan Z., Kirschner J. & Štěpánek J. (eds) (2002) Klíč ke květeně České
republiky [Key to the flora of the Czech Republic]. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Reproduction is the production of offspring that are physically separated from the parental plant. Plants reproduce either by seed (or spores) or vegetatively, while the combination of these two types of reproduction in the same taxon is common. Asexual seed production (apomixis) is not considered as vegetative reproduction. The data were taken from the BiolFlor database (Durka 2002).
Durka W. (2002) Blüten- und Reproduktionsbiologie. – In: Klotz S., Kühn I. & Durka W. (eds), BIOLFLOR – Eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland, Schriftenreihe für Vegetationskunde 38: 133–175.
Diaspore, also called dispersule or propagule, is a generative or vegetative part of the plant body that is dispersed from the parental plant and can produce a new individual. Generative diaspores include spores, seeds and fruits or similar dispersal units (e.g. aggregate fruits in Fragaria, multiple fruits in Morus, gymnosperm cones, epimatium-bearing seed in Taxus, spikelets or their various fragments in Poaceae). If the seed is released from dehiscent fruit or decaying ripe fleshy fruit, both seed and fruit can be considered as diaspores. In plants with indehiscent fruits, only the fruit is considered as a diaspore. A specific category of generative diaspore is tumbleweeds, i.e. mature plant parts including stem branches and large inflorescence (e.g. Crambe tataria and Falcaria vulgaris).
Vegetative diaspores are viable and movable parts of plants that originate above ground or in water and disconnect from the parent plant before sprouting. We did not consider as vegetative diaspores clonal organs connected with the maternal plant until the new plant becomes independent (e.g. stolons in Fragaria) and various types of below-ground organs or shoot bases embedded in soil (e.g. tubers of Helianthus tuberosus or grass tillers). Vegetative diaspores include (i) turions (e.g. Myriophyllum and Utricularia) and similar overwintering structures (detachable buds in Elodea and Groenlandia and shortened shoots of some pondweeds produced by rhizome or stolon, e.g. Potamogeton alpinus); (ii) bulbils and tubers of stem origin (e.g. Allium oleraceum and Dentaria bulbifera) or root origin (Ficaria only); (iii) plantlets born by pseudovivipary (e.g. Poa alpina); (iv) plantlets born from buds on leaves (e.g. Cardamine pratensis); (v) plantlets born on free ends of stolons, detachable before establishing (e.g. Hydrocharis and Jovibarba); (vi) unspecialized fragments of the shoot (e.g. Sedum album and many aquatic plants), shoot tips (e.g. Ceratophyllum demersum) or detachable offsprings born from axillary buds (e.g. Agrostis canina, Arabidopsis halleri and Rorippa amphibia); (vii) budding plants (Lemnaceae only); and (viii) gemmae produced by gametophytes (Trichomanes speciosum only).
Sádlo J., Chytrý M., Pergl J. & Pyšek P. (2018) Plant dispersal strategies: a new classification based on themultiple dispersal modes of individual species. – Preslia 90: 1–22.
Plants use different dispersal modes, also called dispersal syndromes, depending on different dispersal vectors. For example, anemochory is the dispersal by wind, hydrochory by water, epizoochory by attachment to an animal body and endozoochory by animals via ingestion. However, single plant species usually use a combination of several dispersal modes rather than a single mode. Distinct combinations of dispersal modes repeatedly occurring in different plant taxa are called dispersal strategies. Sádlo et al. (2018) distinguished nine dispersal strategies named for the genus names of typical representatives. Taxa of the Czech flora are assigned to individual strategies based on this source.
Categories
Sádlo J., Chytrý M., Pergl J. & Pyšek P. (2018) Plant dispersal strategies: a new classification based on themultiple dispersal modes of individual species. – Preslia 90: 1–22.
Myrmecochorous plants, i.e. taxa dispersed by ants, possess an elaiosome, a nutrient-rich fleshy appendage of seed or fruit. However, in many taxa, the morphological indication or direct evidence of myrmecochory is equivocal. Removal experiments (seeds with and without elaiosome offered to ants) or chemical analysis (different nutrient content between seed and elaiosome; Konečná et al. 2018) would be needed to decide whether the appendage is elaiosome or not. Therefore, more categories than a simple binary distinction between myrmecochorous and non-myrmecochorous are recognized here:
Plant taxa that are often carried by ants to the nest although having no elaiosome (e.g. cheaters in this plant-ant mutualism or plant parts used as a building material for ant hills) are classified as non-myrmecochorous.
The data are based on the literature search and examination of seed samples of the taxa that are reported as myrmecochorous and their closely related congenerics. The list of these taxa with seed images is available at http://botanika.prf.jcu.cz/myrmekochorie/. These taxa were selected from the families represented in the Czech flora that contain at least one taxon reported as myrmecochorous in the literature (Sernander 1906, Hejný et al. 1988 onwards, Fitter & Peat 1994, Klotz et al. 2002, Grime et al. 2007, Kleyer et al. 2008, Servigne 2008, Lengyel et al. 2010, Študent 2012). Such taxa were found in 37 families including Amaryllidaceae, Apiaceae, Apocynaceae, Aristolochiaceae, Asparagaceae, Asteraceae, Boraginaceae, Campanulaceae, Caryophyllaceae, Celastraceae, Colchicaceae, Crassulaceae, Cyperaceae, Dipsacaceae, Euphorbiaceae, Fabaceae, Iridaceae, Juncaceae, Lamiaceae, Liliaceae, Linaceae, Montiaceae, Orobanchaceae, Oxalidaceae, Papaveraceae, Plantaginaceae, Poaceae, Polygalaceae, Polygonaceae, Portulacaceae, Primulaceae, Ranunculaceae, Resedaceae, Rosaceae, Santalaceae, Urticaceae and Violaceae. All the taxa not belonging to these families were classified as non-myrmecochorous (b).
For each of the five categories, a subcategory nv (= non vidimus, i.e. not seen) is used in the taxa for which we found neither information in the literature nor a photograph of a seed, and failed to collect seeds from living plants, but the assignment to the category is likely based on the traits of closely related taxa. For example, we have no data for Centaurea bruguiereana but we classify it as myrmecochorous nv, because all the taxa of Centaurea for which we have data possess an elaiosome.
Konečná M., Štech M. & Lepš J. (2018) Myrmecochory. – www.pladias.cz.
Fitter A. H. & Peat H. J. (1994) The Ecological Flora Database. – Journal of Ecology 82: 415–425.
Grime J. P., Hodgson J. G. & Hunt R. (eds) (2007) Comparative plant ecology: a functional approach to common British species. 2nd edition. – Castlepoint Press, Colvend, Dalbeattie.
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Kleyer M., Bekker R. M., Knevel I. C., Bakker J. P., Thompson K., Sonnenschein M., Poschlod P., van Groenendael J. M., Klimeš L., Klimešová J., Klotz S., Rusch G. M., Hermy M., Adriaens D., Boedeltje G., Bossuyt B., Dannemann A., Endels P., Götzenberger L., Hodgson J. G., Jackel A. K., Kühn I., Kunzmann D., Ozinga W. A., Romermann C., Stadler M., Schlegelmilch J., Steendam H. J., Tackenberg O., Wilmann B., Cornelissen J. H. C., Eriksson O., Garnier E. & Peco B. (2008) The LEDA Traitbase: a database of life-history traits of the Northwest European flora. – Journal of Ecology 96: 1266–1274.
Klotz S., Kühn I. & Durka W. (eds) (2002) BIOLFLOR: eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland. – Schriftenreihe für Vegetationskunde 38: 1–334.
Konečná M., Moos M., Zahradníčková H., Šimek P. & Lepš J. (2018) Tasty rewards for ants: differences in elaiosome and seed metabolite profiles are consistent across species and reflect taxonomic relatedness. – Oecologia 188: 753–764.
Sernander R. (1906) Entwurf einer Monographie der europäischen Myrmekochoren. – Kungliga Svenska Vetenskapsakademiens Handlingar 41: 1–410.
Servigne P. (2008) Etude expérimentale et comparative de la myrmécochorie: le cas des fourmis dispersatrices Lasius niger et Myrmica rubra. – PhD thesis, Université libre de Bruxelles, Bruxelles.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Študent V. (2012) Společné funkční vlastnosti myrmekochorních druhů rostlin České republiky a sezónní a denní dynamika odnosu diaspor všivce lesního (Pedicularis sylvatica) mravenci [Traits of myrmecochorous plants of the Czech Republic and a seasonal and daily seed’s removal dynamics of lousewort (Pedicularis sylvatica) by ants]. – Master thesis, University of South Bohemia, České Budějovice.
Root metamorphoses are modifications of the root that involve the development of different structures for special tasks such as vegetative spread or storage. Types of root metamorphoses are adopted from the BiolFlor database (Krumbiegel 2002).
Categories
Krumbiegel A. (2002) Morphologie der vegetativen Organe (außer Blätter). – In: Klotz S., Kühn I. & DurkaW. (eds), BIOLFLOR: eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland, Schriftenreihe für Vegetationskunde 38: 93–118.
The type of clonal growth is only reported for clonal herbs. Clonal growth is defined here as the growth of the plant body leading to the formation of physically independent asexual offspring. A morphological prerequisite for clonal growth is the formation of adventitious roots on stems or adventitious shoots from root buds that yield (potentially) physically independent individuals (Groff & Kaplan 1988). The types of clonal growth organs are morphological categories that are defined based on three main parameters:
For each taxon, only one type of the clonal growth organ is reported, although some taxa possess several independent types of such organs (Klimešová & Klimeš 2006). The reported type is considered as the most important for the life cycle of the taxon, producing the highest number of offspring or permitting the individual to spread its offspring over large distances. Some of these types are vegetative diaspores, while others are used for local spread but not long-distance dispersal. The clonal growth organs are divided into aboveground and belowground and sorted within each category by their decreasing frequency in the Czech flora.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European plants. – http://clopla.butbn.cas.cz.
Groff P. A. & Kaplan D. R. (1988) The relation of root systems to shoot systems in vascular plants. – Botanical Review 54: 387–422.
This trait is defined only for clonal herbs. Clonality of herbs can be realized by the formation of freely dispersible clonal offspring, i.e. new individuals that are separated from the mother shoots very shortly after their formation and before they develop roots attaching them to the soil. They are dispersed by water or other agents. Typical examples are plantlets, bulbils, turions or stem fragments of aquatic plants. The data reported here are based on individual observations in the CLO-PLA 3.4 database (Klimešová & Klimeš 2006, Klimešová et al. 2017).
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
This trait, defined for herbs, is measured as the number of years from the emergence of the aboveground part of the shoot till its flowering and fruiting (Serebryakov 1952). Based on the analysis of morphological traits, we distinguish shoots with cyclicity of one year (monocyclic) from those that live longer (di- and polycyclic). In plants with sympodial branching, cyclicity refers to all shoots, while in plants with monopodial branching, it refers only to flowering shoots, although flowering and sterile shoots can be present simultaneously. Monocyclic plants usually do not possess a leaf rosette, and all shoots in a population can flower. In contrast, di- and polycyclic shoots possess a basal leaf rosette and shoot populations contain flowering and sterile shoots at the same time.
The data are based on individual observations in the CLO-PLA 3.4 database (Klimešová & Klimeš 2006). If more types are reported for one taxon, the most frequently observed type is given (Klimešová et al. 2017).
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
Serebryakov I. G. (1952) Morfologiya vegetativnykh organov vysshikh rastenii [Morphology of vegetative organs of higher plants]. – Sovetskaya nauka, Moskva.
Branching type is defined for clonal herbs. It determines whether individuals possess two different shoot types (flowering and sterile) or only one shoot type (which can potentially flower). In plants with sympodial branching, all shoots are identical in their construction, replacing each other during ontogeny of the individual; all of them can potentially flower. In contrast, plants with monopodial branching possess two shoot types, one of which never flowers, whereas the flowering shoots arise from axillary buds of the non-flowering shoot. Finally, ferns and lycophytes can possess dichotomous branching that is functionally similar to monopodial branching.
The data reported here are based on individual observations in the CLO-PLA 3.4 database (Klimešová & Klimeš 2006). If more types are reported for one taxon, the most frequently observed type is given here (Klimešová et al. 2017).
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
The presence of the primary root is only defined for herbs. The primary root can be either present for the whole life of a plant or replaced during the ontogeny by adventitious roots. If the primary root is the only root for the whole life of a plant, the plant is not capable of forming adventitious roots on stems; therefore it is not clonal (unless it is able to form adventitious buds on roots; Groff & Kaplan 1988). In contrast, if the primary root is existing only in an early ontogenetic stage and later replaced by adventitious roots formed on belowground parts of the stem, the plant can grow clonally. In older individuals of some taxa that preserve the primary root, this root can split into parts, giving rise to several independent plant individuals. Some taxa only form adventitious roots under specific conditions (soil moisture, root injury or old age).
The data reported here are based on individual observations stored in the CLO-PLA 3.4 database (Klimešová & Klimeš 2006). If more types are reported for one taxon, the most frequently observed type is given (Klimešová et al. 2017).
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
Groff P. A. & Kaplan D. R. (1988) The relation of root systems to shoot systems in vascular plants. – Botanical Review 54: 387–422.
Persistence of the clonal growth organ, defined for clonal herbs, determines the life span of the physical connection between the parent and offspring shoots. Because morphological analysis does not permit the identification of such life span beyond a period of few years, the persistence of the connection is assessed in categories (< 1, 1–2, > 2 years; Klimešová & Klimeš 2006). From those categories, mean values of their ranges (0.5, 1.5 and 4 years) are used, and the final value is the mean of all records for the given taxon and the given type of the clonal growth organ in the CLO-PLA 3.4 database (Klimešová et al. 2017).
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
This trait is only defined for clonal herbs. The number of offspring shoots produced per parent shoot of a clonal herb per year is estimated in categories (< 1, 1, 2–10, > 10; Klimešová & Klimeš 2006), which are represented by the mean values of their ranges (0.5, 1, 6, and 15). The reported value is the mean of these values across all the available measurements for individuals of the given taxon and type of clonal growth organ in the CLO-PLA 3.4 database (Klimešová et al. 2017).
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
The lateral spreading distance by clonal growth is defined for clonal herbs as the distance between parental and offspring shoots. Freely dispersible vegetative diaspores are not considered. Lateral spreading distances were estimated in categories (< 0.01 m, 0.01–0.25 m, > 0.25 m; Klimešová & Klimeš 2006), which are represented by the mean values of their ranges (0.005 m, 0.13 m, 0.5 m). The reported value is the mean of these values across all records for the given taxon and the given type of clonal growth organ in the CLO-PLA 3.4 database (Klimešová et al. 2017).
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
The Clonal index (Johansson et al. 2011) is a measure of taxon’s clonal ability. It is defined for clonal herbs as the sum of the ranks of the four categories of “Number of clonal offspring” (coded as 1, 2, 3, 4) and the three categories of “Lateral spreading distance by clonal growth” (coded as 1, 2, 3) with the presence of freely dispersible vegetative diaspores added as the fourth category (4). The index values range from 2 to 8, with higher values indicating better clonal ability. The index is defined for clonal herbs.
The data reported here are based on the categories of “Number of clonal offspring” and “Lateral spreading distance by clonal growth” aggregated from individual records in the CLO-PLA 3.4 database (Klimešová & Klimeš 2006, Klimešová et al. 2017).
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
Johansson V. A., Cousins S. A. O. & Eriksson O. (2011) Remnant populations and plant functional traits in abandoned semi-natural grasslands. – Folia Geobotanica 46: 165–179.
In contrast to stem buds, which are always located in a leaf axil, adventitious buds on roots (or hypocotyl) are not located at specific positions of the roots. They occur only in some root types or may only appear after the individual is injured. Root buds are found in approximately 10% of species in the Czech flora. Adventitious buds may be located on the hypocotyl, on the primary root, or on lateral roots. We list the hypocotyl only if the roots are found exclusively on it. We report the primary root if the buds are never formed on lateral roots, and we report the lateral roots if root buds are potentially found on all three types of bud-bearing organs. The data were taken from Bartušková et al. (2017).
Bartušková A., Malíková L. & Klimešová J. (2017) Checklist of root-sprouters in the Czech flora: mapping the gaps in our knowledge. – Folia Geobotanica 52: 337–343.
Adventitious buds on roots or hypocotyl occur in only about 10% species of the Czech flora and have different roles in individual taxa. In some taxa, they occur only after the plant is injured and play a role in regeneration after such injury (regenerative role). In other taxa, these buds are formed without such an external stimulus. Consequently, they increase the number of shoots and thereby the number of offspring, both clonally and by seed. In most taxa, such buds occur only in some individuals and are not necessary for completing the life cycle; we denote them as additive (or accessory) root buds. In a few taxa, the formation of root buds is necessary for completing the life cycle and flowering. They develop in all individuals, and survival/flowering or overwintering of an individual is dependent on them (Rauh 1937, Klimešová 2007). The data were taken from Bartušková et al. (2017).
Bartušková A., Malíková L. & Klimešová J. (2017) Checklist of root-sprouters in the Czech flora: mapping the gaps in our knowledge. – Folia Geobotanica 52: 337–343.
Klimešová J. (2007) Root-sprouting in myco-heterotrophic plants: prepackaged symbioses or overcoming meristem limitation? – New Phytologist 173: 8–10.
Rauh W. (1937) Die Bildung von Hypocotyl- und Wurzelsprossen und ihre Bedeutung für die Wuchsformen der Pflanzen. – Acta Nova Leopoldina 4/24: 395–555.
Bud bank denotes all inactive (dormant) buds on the plant body that can give rise to new shoots, including both shoot buds and root buds (Klimešová & Klimeš 2007). The most important part of the bud bank is located at the soil surface or belowground, out of the reach of disturbance or seasonal frost or drought (Raunkiaer 1934). Consequently, only data on buds located at the soil surface or in the soil are reported here.
The number of buds on plant organs located at different soil depths was assessed according to morphological characters (Klimešová & Klimeš 2007). The assessment was based on the assumption that each leaf (or leaf scale) axil contains a bud. Assessment of bud numbers in individual plants was done in three categories (0, 0–10, > 10 buds per shoot; Klimešová & Klimeš 2006). These categories were respectively represented by values of 0, 5, 15 buds per shoot. The value for the taxon was calculated as the mean of these values across the individuals of this taxon and particular soil depth as reported in the CLO-PLA 3.4 database (Klimešová et al. 2017). The size of the belowground bud bank was determined as the sum of bud numbers per shoot summed over the soil profile. The depth of the belowground bud bank was determined as the average depth of the buds in the soil. In addition to stem-derived buds, around 10% of taxa in the Czech flora possess the ability to form adventitious buds on the root or hypocotyl (here collectively called root buds). As root buds cannot be counted (they are formed freely along the root), 15 buds were arbitrarily added per each 10 cm of depth for categories that include root buds. All the bud-bank characteristics are given for stem-derived buds only (root buds excluded) and all the buds (root buds included):
Klimešová J., Danihelka J., Chrtek J., de Bello F. & Herben T. (2017) CLO-PLA: a database of clonal and budbank
traits of the Central European flora. – Ecology 98: 1179.
Klimešová J. & Klimeš L. (2006) CLO-PLA3: a database of clonal growth architecture of Central-European
plants. – http://clopla.butbn.cas.cz.
Klimešová J. & Klimeš L. (2007) Bud banks and their role in vegetative regeneration: a literature review and proposal for simple classification and assessment. – Perspectives in Plant Ecology, Evolution and Systematics 8: 115–129.
Raunkiaer C. (1934) The life forms of plants and statistical plant geography. – Clarendon Press, Oxford.
Plant parasitism is based on either of two mechanisms. The first group of parasitic plants involves those parasitizing directly on another plant. These plants are called haustorial parasites. They take resources from the host’s vascular bundles using a specialized organ, the haustorium. The second group comprises mycoheterotrophic plants, which parasitize fungi via mycorrhizal interaction and gain organic carbon from them.
Plants in both groups display variable dependence on their host organism. The haustorial parasites include two distinct functional groups: green hemiparasites and holoparasites. Green hemiparasites are partial parasites that retain photosynthetic ability but obtain all mineral resources and a part of the organic carbon from the host. Holoparasites are non-green full parasites unable to photosynthesize. Location of the haustorial attachment to the host (root or stem) is another essential functional trait. The distinction between partial and full parasitism in haustorial parasites may not be straightforward. In the Czech flora, it is nevertheless possible to distinguish between stem hemi- and holoparasites, which are difficult to separate on the global scale (Těšitel 2016). Consequently, we use a traditional classification here and classify as holoparasites those plants that are in adulthood mostly without chlorophyll, even though some of them might have some chlorophyll and perform residual photosynthesis (e.g. Cuscuta).
In mycoheterotrophic plants, there is a continuum from initial mycoheterotrophs through partial mycoheterotrophs to full mycoheterotrophs. In the initial mycoheterotrophs, only initial stages, i.e. gametophytes or seedlings, are dependent on the fungus, whereas adult plants are autotrophic, while still depending on mycorrhizal symbiosis as a source of water and mineral nutrients. In the partial mycoheterotrophs, photosynthesizing adults obtain from their mycorrhizal fungi not only water and mineral nutrients but also different amounts of organic carbon. The full mycoheterotrophs lost their chlorophyll and are thus fully parasitic. In some partial mycoheterotrophs (e.g. the genus Cephalanthera), chlorotic individuals can be found, which lack chlorophyll and fully depend on their hosts.
Classification of haustorial parasites follows Těšitel (2016) with a further distinction of stem hemi- and holoparasites, and identification of mycoheterotrophs follows Merckx (2012).
Těšitel J., Těšitelová T., Blažek P. & Lepš J. (2016) Parasitism and mycoheterotrophy. – www.pladias.cz.
Těšitel J. (2016) Functional biology of parasitic plants: a review. – Plant Ecology and Evolution 149: 5–20.
Merckx V. S. F. T. (2012) Mycoheterotrophy: the biology of plants living on fungi. – Springer, Berlin.
Carnivorous plants attract, trap and kill their prey, animals (mainly insects and small crustaceans) and protozoans, and subsequently absorb the nutrients from their dead bodies.
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Plants are classified into those without symbiotic nitrogen fixers and those that form a symbiosis with nitrogen-fixing bacteria. The latter are further divided into those forming a symbiosis with rhizobia (e.g. Allorhizobium, Bradyrhizobium, Mesorhizobium, Rhizobium and Sinorhizobium) and those forming the actinorhizal symbiosis with the genus Frankia, the latter called actinorhizal plants (Bond 1983, Pawlowski & Sprent 2007, Sprent 2008, Benson 2016).
In the Czech flora, the rhizobial group is represented by virtually all legumes (family Fabaceae). Exceptions are three non-native cultivated woody species (Cercis siliquastrum, Gleditsia triacanthos, Gymnocladus dioicus) that do not nodulate (Tedersoo et al. 2018), which is generally considered as evidence for the absence of symbiosis. However, some studies suggest that functional nitrogen-fixing symbiosis may exist even without visible nodules (Bryan et al. 1996). Roots of Gleditsia triacanthos were recorded to contain bacterial structures similar to those in nodules with rhizobia, as well as the presence of nitrogenase (Faria et al. 2002). These genera also contain genes probably related to nodule formation, although their exact function is unclear (Graves et al. 1999). Because convincing evidence of nitrogen fixation in these species is missing, we consider them non-nitrogen-fixing for the time being.
Symbiosis with rhizobia was found in several other families (Tedersoo et al. 2018). Of these, the Czech flora includes only casually introduced Tribulus terrestris (Zygophyllaceae), in which a parallel infection with cyanobacteria was described (Sabet 1946, Mahmood & Athar 1998).
The actinorhizal group is represented in the Czech flora mainly by alder species (Alnus spp.) and also by cultivated species in the family Elaeagnaceae – Elaeagnus spp. and Hippophaë rhamnoides (Bond 1983, Benson 2016).
Blažek P. & Lepš J. (2016) Symbiotic nitrogen fixation. – www.pladias.cz.
Benson D. R. (2016) Frankia & actinorhizal plants. – https://frankia.mcb.uconn.edu/ (accessed on 1 Feb 2021).
Bond G. (1983) Taxonomy and distribution of non-legume nitrogen-fixing systems. – In: Gordon J. C. & Wheeler C. T. (eds), Biological nitrogen fixation in forests: foundations and applications, p. 55–87, Martinus Nijhoff/Dr W. Junk Publ., The Hague.
Bryan J. A., Berlyn G. P. & Gordon J. C. (1996) Toward a new concept of the evolution of symbiotic nitrogen fixation in the Leguminosae. – Plant and Soil 186: 151–159.
de Faria S. M., Olivares F. L. & Xavier R. P. (2002) Nodule-structure in the roots of Gleditsia spp. a non-nodulating legume genus. – In: Pedrosa F. O., Hungria M., Yates G. & Newton W. E. (eds), Nitrogen fixation: from molecules to crop productivity. Current plant science and biotechnology in agriculture, vol 38. Springer, Dordrecht, p. 337.
Graves W. R., Foster C. M., Rosin F. M. & Schrader J. A. (1999) Two early nodulation genes are not markers for the capacity of leguminous nursery crops to form root nodules. – Journal of Environmental Horticulture 17: 126–129.
Mahmood A. & Athar M. (1998) Cyanobacterial root nodules in Tribulus terrestris L. (Zygophyllaceae). – In: Malik K. A. & Sajjad Mirza M. & Ladha J. K. (eds), Nitrogen fixation with non-legumes, Springer, Dordrecht, p. 345–350.
Pawlowski K. & Sprent J. I. (2007) Comparison between actinorhizal and legume symbioses. – In: Pawlowski K. & Newton W. E. (eds), Nitrogen-fixing actinorhizal symbioses, Springer, Dordrecht, p. 261–288.
Sabet Y. S. (1946) Bacterial root nodules in the Zygophyllaceae. – Nature 157: 656–657.
Sprent J. I. (2008) Evolution and diversity of legume symbiosis. – In: Dilworth M. J., James E. K., Sprent J. I. & Newton W. E. (eds), Nitrogen-fixing leguminous symbioses, Springer, Dordrecht, p. 1–21.
Tedersoo L., Laanisto L., Rahimlou S., Toussaint A., Hallikma T. & Pärtel M. (2018) Global database of plants with root-symbiotic nitrogen fixation: NodDB. – Journal of Vegetation Science 29: 560–568.
Chromosome number is the somatic number of chromosomes in the zygotic stage, i.e. without possible endopolyploidy of somatic tissues. If different chromosome numbers are known for a taxon, the database contains primarily the number reported from the Czech Republic or the number that is the most common in this country or can be expected to be the most common based on the data from neighbouring countries. Other existing and less common chromosome numbers are reported in brackets. The survey does not take into account odd chromosome numbers of individual, aneuploid, euploid, haploid or autopolyploid plants, which may rarely originate in natural or experimental populations. It also disregards the numbers reported in early studies or from geographically distant areas for which the taxonomic identity with Czech plants is unclear.
The data compilation is based mainly on the Flora of the Czech Republic (vols. 1–8; Hejný et al. 1988–1992, Slavík et al. 1997–2004, Štěpánková et al. 2010) and the Chromosome Count Database (Rice et al. 2015; http://ccdb.tau.ac.il/). If only information about ploidy level is available from flow cytometry measurements, but no chromosome number is known, the number typical of the given ploidy in closely related taxa is indicated.
Šmarda P. (2018) Chromosome number (2n). – www.pladias.cz.
Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds) (1988) Květena České socialistické republiky [Flora of the Czech Socialist Republic]. Vol. 1. – Academia, Praha.
Hejný S., Slavík B., Hrouda L. & Skalický V. (eds) (1990) Květena České republiky [Flora of the Czech Republic]. Vol. 2. – Academia, Praha.
Hejný S., Slavík B., Kirschner J. & Křísa B. (eds) (1992) Květena České republiky [Flora of the Czech Republic]. Vol. 3. – Academia, Praha.
Rice A., Glick L., Abadi S., Einhorn M., Kopelman N. M., Salman-Minkov A., Mayzel J., Chay O. & Mayrose I. (2015) The chromosome counts database (CCDB): a community resource of plant chromosome numbers. – New Phytologist 206: 19–26.
Slavík B., Chrtek J. jun. & Štěpánková J. (eds) (2000) Květena České republiky [Flora of the Czech Republic]. Vol. 6. – Academia, Praha.
Slavík B., Chrtek J. jun. & Tomšovic P. (eds) (1997) Květena České republiky [Flora of the Czech Republic]. Vol. 5. – Academia, Praha.
Slavík B., Smejkal M., Dvořáková M. & Grulich V. (eds) (1995) Květena České republiky [Flora of the Czech Republic]. Vol. 4. – Academia, Praha.
Slavík B., Štěpánková J. & Štěpánek J. (eds) (2004) Květena České republiky [Flora of the Czech Republic]. Vol. 7. – Academia, Praha.
Štěpánková J., Chrtek J. jun. & Kaplan Z. (eds) (2010) Květena České republiky [Flora of the Czech Republic]. Vol. 8. – Academia, Praha.
Ploidy level is the number of somatic chromosomal sets in the zygotic stage, i.e. without the possible endopolyploidy of somatic tissues. Ploidy level determines the minimum copy number of most genes, influences minimal cell size and other morphological and ecological properties of the taxon (Stebbins 1950, Levin 2002, Tate et al. 2005). The data presented here are based on the traits Chromosome number and 2C genome size, and to a lesser extent also on a literature search of flow cytometry studies related to the area of the Czech Republic. The reported values are especially those reported from the Czech Republic or those ploidy levels that are most frequent in this country or at least are assumed to be the most frequent based on the data from neighbouring countries. The other existing (minor) ploidy levels (cytotypes) that are documented from the Czech Republic or that may be expected to occur here based on the records from neighbouring countries are indicated in brackets. The survey does not take into account the observations of individual haploid or autopolyploid plants which may rarely originate in natural or experimental populations. It also disregards the ploidy levels derived from the numbers of chromosomes reported in early studies or from geographically distant areas, where taxonomic identity with Czech plants is uncertain.
The size of one chromosomal set (x) or the “base chromosome number” for calculating the ploidy level is derived here from the lowest chromosome number known in the given genus or the group of closely related genera (e.g. Raven 1975). This minimum chromosome number is generally considered to correspond to diploids (i.e., two chromosomal sets). A taxon is considered as polyploid whenever its chromosome number and genome size are jointly ± doubled (or otherwise multiplied) compared to the diploid taxa of related genera. It may therefore sometimes happen that no diploid taxa are known in some genera. The absence of diploids in a given genus may result from (i) the lack of karyological data, (ii) the extinction of the diploid relative(s), or (iii) a polyploidy event that predated the origin of the whole genus, with the current genomes still showing little signs of backward “diploidization”. The joint usage of the chromosome number and genome size enables estimation of ploidy levels also for the taxa with holocentric chromosomes (Cyperaceae, Drosera, Juncaceae, Cuscuta sect. Cuscuta and C. sect. Gramica). In these taxa, the chromosome number does not need to be positively correlated with the ploidy level due to possible chromosomal fissions and fusions (agmatoploidy and symploidy, respectively; Bureš et al. 2013). To handle the chromosomal fusions in Luzula, chromosome size categories as defined by Nordeskiöld (1951) were further considered to estimate the actual ploidy level. Ploidy level estimates in highly polyploid genomes of Viola follow Marcussen et al. (2015).
Šmarda P. (2018) Ploidy level (x). – www.pladias.cz.
Bureš P., Zedek F. & Marková M. (2013) Holocentric chromosomes. – In: Leitch I. J., Greilhuber J., Wendel J. & Wendel J. (eds), Plant genome diversity. Vol. 2. Physical structure, behaviour and evolution of plant genomes, p. 187–208, Springer, Wien.
Dar T.-U.-H. & Rehman R.-U. (2017) Polyploidy: recent trends and future perspectives. – Springer, New Delhi.
Levin D. A. (2002) The role of chromosomal change in plant evolution. – Oxford University Press, Oxford.
Nordenskiöld H. (1951) Cytotaxonomical studies in the genus Luzula. – Hereditas 37: 325–355.
Raven P. H. (1975) The bases of angiosperm phylogeny: cytology. – Annals of the Missouri Botanical Garden
62: 724–764.
Stebbins G. L. (1950) Variation and evolution in plants. – Columbia University Press, New York.
Tate J. A., Soltis D. E. & Soltis P. S. (2005) Polyploidy in plants. – In: Gregory R. T. (ed.), The evolution of the
genome, p. 371–426, Elsevier, Amsterdam.
2C genome size is the somatic nuclear DNA content in a zygotic cell measured in megabase pairs (Mbp). This measure can vary among taxa due to both polyploidy and the variability in the content of non-coding DNA (Leitch & Greilhuber 2013). Genome size influences minimum cell size, duration of the cell cycle and cell division, and nutrient requirements. Therefore, it may have a considerable influence on ecological strategies of plants (Bennett 1987, Veselý et al. 2012, Greilhuber & Leitch 2013). Most values were measured in plants collected in the Czech Republic (Šmarda et al. 2019). The data always refer to the dominant chromosome number and the dominant ploidy level of the given taxon.
Šmarda P., Knápek O., Březinová A., Horová L., Grulich V., Danihelka J., Veselý P., Šmerda J., Rotreklová O. & Bureš P. (2019) Genome sizes and genomic guanine+cytosine (GC) contents of the Czech vascular flora with new estimates for 1700 species. – Preslia 91: 117–142.
Bennett M. D. (1987) Variation in genomic form in plants and its ecological implications. – New Phytologist 106 (Suppl.): 177–200.
Greilhuber J. & Leitch I. J. (2013) Genome size and the phenotype. – In: Leitch I. J., Greilhuber J., Doležel J. & Wendel J. (eds), Plant genome diversity. Vol. 2. Physical structure, behaviour and evolution of plant genomes, p. 323–344, Springer, Wien.
Leitch I. J. & Greilhuber J. (2013) Genome size diversity and evolution in land plants. – In: Leitch I. J., Greilhuber J., Doležel J. & Wendel J. (eds), Plant genome diversity. Vol. 2. Physical structure, behaviour and evolution of plant genomes, p. 307–322, Springer, Wien.
Veselý P., Bureš P., Šmarda P. & Pavlíček T. (2012) Genome size and DNA base composition of geophytes: the mirror of phenology and ecology? – Annals of Botany 109: 65–75.
1Cx monoploid genome size is the amount of DNA contained in one set of chromosomes measured in megabase pairs (Mbp). The values were obtained for each taxon by dividing its 2C genome size by the respective ploidy level (Greilhuber et al. 2005). Differences in 1Cx values among taxa are therefore virtually free of the polyploidy effect (i.e. only due to amplification of non-coding DNA). However, the 1Cx values in polyploids are usually slightly smaller due to the increased tendency to eliminate the duplicated, redundant DNA (Leitch & Bennett 2004). Because the 1Cx values tend to be similar in related taxa, they can be used to roughly estimate the 2C genome size in related taxa for which only the ploidy level is known so far. Conversely, they can be used to estimate the ploidy levels based on the known 2C genome size. The data were taken from Šmarda et al. (2019).
Šmarda P., Knápek O., Březinová A., Horová L., Grulich V., Danihelka J., Veselý P., Šmerda J., Rotreklová O. & Bureš P. (2019) Genome sizes and genomic guanine+cytosine (GC) contents of the Czech vascular flora with new estimates for 1700 species. – Preslia 91: 117–142.
Greilhuber J., Doležel J., Lysák M. A. & Bennett M. D. (2005) The origin, evolution and proposed stabilization of the terms ‘genome size’ and ‘C-value’ to describe nuclear DNA content. – Annals of Botany 95: 255–260.
Leitch I. J. & Bennett M. D. (2004) Genome downsizing in polyploid plants. – Biological Journal of the Linnean Society 82: 651–663.
Genomic GC content is the percentage of guanine and cytosine bases in nuclear DNA. It influences the thermal stability of DNA, packing of condensed DNA within the nucleus, the energetic cost of DNA synthesis or cell sensitivity to desiccation (Šmarda & Bureš 2012, Šmarda et al. 2014). For the vast majority of taxa, these data were measured in plants collected in the Czech Republic (Šmarda et al. 2019). The data always refer to the dominant chromosome number and dominant ploidy of the given taxon. Differences up to 1% in closely related taxa or up to 2% in unrelated taxa may be considered insignificant because of possible method errors (Šmarda et al. 2012).
Šmarda P., Knápek O., Březinová A., Horová L., Grulich V., Danihelka J., Veselý P., Šmerda J., Rotreklová O. & Bureš P. (2019) Genome sizes and genomic guanine+cytosine (GC) contents of the Czech vascular flora with new estimates for 1700 species. – Preslia 91: 117–142.
Šmarda P. & Bureš P. (2012) The variation of base composition in plant genomes. – In: Wendel J., Greilhuber J., Doležel J. & Leitch I. J. (eds), Plant genome diversity. Vol. 1. Plant genomes, their residents, and their evolutionary dynamics, p. 209–235, Springer, Heidelberg.
Šmarda P., Bureš P., Šmerda J. & Horová L. (2012) Measurements of genomic GC content in plant genomes with flow cytometry: a test for reliability. – New Phytologist 193: 513–521.
Šmarda P., Bureš P., Horová L., Leitch I. J., Mucina L., Pacini E., Tichý L., Grulich V. & Rotreklová O. (2014) Ecological and evolutionary significance of genomic GC content diversity in monocots. – Proceedings of the National Academy of Sciences of the USA 111: E4096–E4102.
Taxa are classified according to whether they are native or alien to the Czech Republic. Following the definitions used in invasion ecology, native taxa are those that have evolved in the area of the Czech Republic or immigrated there without human assistance from the area where they had evolved. Alien taxa are those whose presence is a result of intentional or unintentional introduction by human activity and can be divided based on their residence time. The alien taxa are divided based on their residence time into archaeophytes and neophytes. Archaeophytes are taxa occurring in the wild that were introduced between the beginning of Neolithic agriculture and the year 1500, i.e. the beginning of intercontinental overseas trade after the discovery of the Americas. Neophytes are taxa occurring in the wild that were introduced after 1500 (see Richardson et al. 2000 for detailed definitions). Some taxa introduced in the Late Middle Ages or Early Modern Period, but with no exact information on the introduction date, were assigned to a joint category of Archaeophyte/neophyte. Additionally, some frequently cultivated taxa that are not known to have escaped from cultivation are listed as a separate category Cultivated. Category Lack of evidence of occurrence in the wild includes taxa for which spontaneous occurrence in the wild is doubtful. Taxa assigned to the category Absent in Czechia are not sufficiently supported by reliable records or occurred just once and disappeared.
The data included in the database follow the third edition of the Catalogue of alien plants of the Czech Republic (Pyšek et al. 2022 and references related to individual taxa therein).
Pyšek P., Sádlo J., Chrtek J. Jr., Chytrý M., Kaplan Z., Pergl J., Pokorná A., Axmanová I., Čuda J., Doležal J., Dřevojan P., Hejda M., Kočár P., Kortz A., Lososová Z., Lustyk P., Skálová H., Štajerová K., Večeřa M., Vítková M., Wild J. & Danihelka J. (2022) Catalogue of alien plants of the Czech Republic (3rd edition): species richness, status, distributions, habitats, regional invasion levels, introduction pathways and impacts. – Preslia 94: 447–577.
Richardson D. M., Pyšek P., Rejmánek M., Barbour M. G., Panetta F. D. & West C. J. (2000) Naturalization and invasion of alien plants: concepts and definitions. – Diversity and Distributions 6: 93–107.
Indicator value for light is expressed on an ordinal scale from 1 to 9 defined by Ellenberg et al. (1991). The values for individual taxa have been modified and extended for the Czech flora by Chytrý et al. (2018). Values with “x” indicate generalists, i.e. taxa with broad ecological range with respect to light. Indicator values for trees relate to juvenile individuals in herb and shrub layer.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018) Ellenberg-type indicator values for the Czech flora. – Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991) Zeigerwerte von Pflanzen in Mitteleuropa. – Scripta Geobotanica 18: 1–248.
Indicator value for temperature is expressed on an ordinal scale from 1 to 9 defined by Ellenberg et al. (1991). The values for individual taxa have been modified and extended for the Czech flora by Chytrý et al. (2018). Values with “x” indicate generalists, i.e. taxa with broad ecological range with respect to temperature.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018) Ellenberg-type indicator values for the Czech flora. – Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991) Zeigerwerte von Pflanzen in Mitteleuropa. – Scripta Geobotanica 18: 1–248.
Indicator value for moisture is expressed on an ordinal scale from 1 to 12 defined by Ellenberg et al. (1991). The values for individual taxa have been modified and extended for the Czech flora by Chytrý et al. (2018). Values with “x” indicate generalists, i.e. taxa with broad ecological range with respect to moisture.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018) Ellenberg-type indicator values for the Czech flora. – Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991) Zeigerwerte von Pflanzen in Mitteleuropa. – Scripta Geobotanica 18: 1–248.
Indicator value for soil or water reaction is expressed on an ordinal scale from 1 to 9 defined by Ellenberg et al. (1991). The values for individual taxa have been modified and extended for the Czech flora by Chytrý et al. (2018). Values with “x” indicate generalists, i.e. taxa with broad ecological range with respect to the reaction. In acidic environments, the value can be considered as a proxy for pH, while in near-neutral or alkaline environments it is more a proxy for calcium concentration.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018) Ellenberg-type indicator values for the Czech flora. – Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991) Zeigerwerte von Pflanzen in Mitteleuropa. – Scripta Geobotanica 18: 1–248.
Indicator value for nutrients is expressed on an ordinal scale from 1 to 9 defined by Ellenberg et al. (1991). The values for individual taxa have been modified and extended for the Czech flora by Chytrý et al. (2018). Values with “x” indicate generalists, i.e. taxa with broad ecological range with respect to nutrient availability. The value is a proxy for availability of nitrogen or phosphorus and to some extent also a proxy for site primary productivity.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018) Ellenberg-type indicator values for the Czech flora. – Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991) Zeigerwerte von Pflanzen in Mitteleuropa. – Scripta Geobotanica 18: 1–248.
Indicator value for salinity is expressed on an ordinal scale from 0 to 9 defined by Ellenberg et al. (1991). The values for individual taxa have been modified and extended for the Czech flora by Chytrý et al. (2018). It is a proxy for concentration in the environment of soluble salts, including sulphates, chlorides and carbonates of sodium, potassium, calcium and magnesium.
Chytrý M., Tichý L., Dřevojan P., Sádlo J. & Zelený D. (2018) Ellenberg-type indicator values for the Czech flora. – Preslia 90: 83–103.
Ellenberg H., Weber H. E., Düll R., Wirth V., Werner W. & Paulißen D. (1991) Zeigerwerte von Pflanzen in Mitteleuropa. – Scripta Geobotanica 18: 1–248.
Indicator values for disturbance express relationships of common taxa of the Czech flora separately to the frequency and severity of disturbance. Individual disturbance agents are not distinguished, but a wide range of factors is considered including logging, cutting, mowing, herbivory, trampling, damage by herbicides, burning, wind-throws, soil erosion, ploughing, hoeing or burrowing, wave and current action, and flooding. There are three types of indicator values for disturbance:
Each of these three types of indicator values is provided separately for the whole-community disturbance events and for smaller disturbance events that affect the herb layer but not the tree layer in forests. Both indicator values are identical for taxa of open habitats.
The following indicator values for disturbance are provided:
Indicator values for disturbance were calculated by Herben et al. (2016) based on an analysis of a stratified subset of 30,115 vegetation plots from the Czech National Phytosociological Database (Chytrý & Rafajová 2003). Only taxa occurring in at least 20 plots were used. These plots were classified by an expert system into 39 phytosociological vegetation classes as defined in Vegetation of the Czech Republic (Chytrý 2007–2013). For each of these classes, the mean frequency and severity of disturbance were assessed based on field observations. The disturbance indicator value for each taxon was calculated as the average disturbance frequency or severity weighted by the frequency of occurrence of that taxon in the plots assigned to these vegetation classes. The structure-based disturbance indicator values were calculated based on the variation in plant height at maturity and variation in summed covers of all taxa recorded in the vegetation plots where the target taxon occurs.
Herben T., Chytrý M. & Klimešová J. (2016) A quest for species-level indicator values for disturbance. – Journal of Vegetation Science 27: 628–636.
Chytrý M. (ed.) (2007–2013) Vegetace České republiky 1–4 [Vegetation of the Czech Republic 1–4]. – Academia, Praha.
Chytrý M. & Rafajová M. (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. – Preslia 75: 1–15.
Data on taxon occurrence in habitats of the Czech Republic are based on the analysis of vegetation plots from the Czech National Phytosociological Database (Chytrý & Rafajová 2003) and its expert revision and completion based on the literature and field experience, especially for rare and taxonomically problematic taxa. The classification recognizes 88 basic habitats aggregated to 13 broader habitats that are defined by Sádlo et al. (2007: their Appendix 1):
Taxon occurrence in each habitat is assessed on a four-degree scale:
Sádlo J., Chytrý M. & Pyšek P. (2007) Regional species pools of vascular plants in habitats of the Czech Republic. – Preslia 79: 303–321.
Chytrý M. & Rafajová M. (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. – Preslia 75: 1–15.
The affinity of taxa to the forest environment is assessed using the categories of the German national list of forest taxa (Schmidt et al. 2011). Each taxon is assessed separately for the region of Thermophyticum (lowlands with thermophilous and drought-adapted flora) and merged regions of Mesophyticum and Oreophyticum (mid-elevations and mountains with mesophilous and mountain flora; Skalický 1988). The compilation was based on the list of regional species pools of Czech habitats (Sádlo et al. 2007), expert knowledge and various literature sources. It has been integrated into the European forest plant species list (Heinken et al. 2019).
Categories
Dřevojan P., Chytrý M., Sádlo J. & Pyšek P. (2016) Affinity to the forest environment. – www.pladias.cz.
Heinken T., Diekmann M., Liira J., Orczewska A., Brunet J., Chytrý M., Chabrerie O., de Frenne P., Decoq G.,
Dřevojan P., Dzwonko Z., Ewald J., Feilberg J., Graae B. J., Grytnes J. A., Hermy M., Kriebitzsch W.-U.,
Laivins M., Lindmo S., Marage D., Marozas V., Meirland A., Niemeyer T., Paal J., Pyšek P., Roosaluste E.,
Sádlo J., Schaminée J., Schmidt M., Tyler T., Verheyen K. & Wulf M. (2019) European forest plant species
list. – Fighshare, https://doi.org/10.6084/m9.figshare.8095217.v1.
Sádlo J., Chytrý M. & Pyšek P. (2007) Regional species pools of vascular plants in habitats of the Czech Republic. – Preslia 79: 303–321.
Schmidt M., KriebitzschW.-U. & Ewald J. (eds) (2011) Waldartenlisten der Farn- und Blütenpflanzen, Moose und Flechten Deutschlands. – BfN-Skripten 299: 1–111.
Skalický V. (1988) Regionálně fytogeografické členění [Regional phytogeographic division]. – In: Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds), Květena České socialistické republiky [Flora of the Czech Socialist Republic] 1: 103–121, Academia, Praha.
Diagnostic taxa are characterized by a concentration of their occurrence in the stands belonging to the target vegetation unit while being rare or absent in other vegetation units. They were determined based on the calculation of fidelity of each taxon to a group of vegetation plots representing the target vegetation unit in a geographically and ecologically stratified selection of plots of all vegetation types from the Czech National Phytosociological Database (Chytrý & Rafajová 2003). Fidelity was measured using the phi coefficient of association after the sizes of plot groups were virtually standardized to 1% of the total size of the data set following Tichý & Chytrý (2006). The taxa with a value of phi higher than 0.25 and significant concentration in the vegetation unit according to Fisher’s exact test (P < 0.001) were considered as diagnostic taxa. The data on the diagnostic status of taxa for individual phytosociological classes, alliances or associations were taken from the monograph Vegetation of the Czech Republic (Chytrý 2007–2013). The numbers of vegetation plots used for the calculations are given in respective volumes of this monograph.
Chytrý M. (ed.) (2007–2013) Vegetace České republiky 1–4 [Vegetation of the Czech Republic 1–4]. – Academia, Praha.
Chytrý M. & Rafajová M. (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. – Preslia 75: 1–15.
Tichý L. & Chytrý M. (2006) Statistical determination of diagnostic species for site groups of unequal size. – Journal of Vegetation Science 17: 809–818.
Constant taxa are characterized by frequent occurrences in stands belonging to the target vegetation unit, but unlike diagnostic taxa, they can also commonly occur in other vegetation units. They were determined based on the calculation of percentage frequency (constancy) of each taxon in a group of vegetation plots representing the target vegetation unit in a geographically and ecologically stratified selection of plots of all vegetation types extracted from the Czech National Phytosociological Database (Chytrý & Rafajová 2003). The taxa with an occurrence frequency in the vegetation unit higher than 40% were considered as constant taxa. The data on the constant status of taxa for individual phytosociological classes, alliances or associations were taken from the monograph Vegetation of the Czech Republic (Chytrý 2007–2013). The numbers of vegetation plots used for the calculations are given in respective volumes of this monograph.
Chytrý M. (ed.) (2007–2013) Vegetace České republiky 1–4 [Vegetation of the Czech Republic 1–4]. – Academia, Praha.
Chytrý M. & Rafajová M. (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. – Preslia 75: 1–15.
The degree of ecological specialization for individual taxa is estimated based on their co-occurrence with other taxa. The underlying assumption is that variation in the composition of co-occurring taxa indicates the range of habitat conditions suitable to this taxon (Fridley et al. 2007). A taxon repeatedly co-occurring with a similar set of taxa across different sites is more likely to be a specialist with a preference for a specific habitat. Conversely, a taxon co-occurring with various taxa across different sites is more likely to be a generalist tolerating a wide range of habitats. The ecological specialization index (ESI) of a taxon is inversely related to beta diversity calculated for the set of sites at which this taxon occurs.
The ecological specialization indices were calculated based on the vegetation plots from the Czech National Phytosociological Database (Chytrý & Rafajová 2003). Three vegetation datasets were selected from a geographically stratified subset of plots from the database: (i) a dataset including all the vegetation types (30,115 plots, 1935 taxa), (ii) a dataset including only non-forest vegetation (24,712 plots, 1875 taxa) and (iii) a dataset including only forest vegetation (5403 plots, 1264 taxa). Whittaker’s multiplicative measure of beta diversity (Whittaker 1960) rarefied to 10 vegetation plots randomly selected from a subset of plots containing the target taxon (β10) was computed for each taxon (Zelený 2009). Outlier plots with very different species composition were removed from the subset before rarefaction, following a recommendation of Botta-Dukát (2012). Because the calculated value of beta diversity decreases with increasing value of taxon specialization, the value of ESI was calculated as ESI = 10 – β10. This value theoretically ranges from 0 to 9, with high values indicating specialists and low values indicating generalists.
Each ESI value is accompanied by a taxon weight, which represents the total number of plots in which this taxon occurs within a particular dataset. The weights can be used as a measure of the reliability of the calculated ESI value for given taxon, which increases with increasing frequency of the taxon in the dataset. Minimum weight is 10, corresponding to the minimum number of occurrences for which ESI was calculated. The theoretical maximum weight is the number of plots in the given dataset.
The following specialization indices and corresponding taxon weights are available (with ranges of values in brackets):
Zelený D. & Chytrý M. (2019) Ecological Specialization Indices for species of the Czech flora. – Preslia 91: 93–116.
Botta-Dukát Z. (2012) Co-occurrence-based measure of species’ habitat specialization: robust, unbiased estimation in saturated communities. – Journal of Vegetation Science 23: 201–207.
Chytrý M. & Rafajová M. (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. – Preslia 75: 1–15.
Fridley J. D., Vandermast D. B., Kuppinger D. M., Manthey M. & Peet R. K. (2007) Co-occurrence based assessment of habitat generalists and specialists: a new approach for the measurement of niche width. – Journal of Ecology 95: 707–722.
Whittaker R. H. (1960) Vegetation of the Siskiyou Mountains, Oregon and California. – Ecological Monographs 30: 279–338.
Zelený D. (2009) Co-occurrence based assessment of species habitat specialization is affected by the size of
species pool: reply to Fridley et al. (2007). – Journal of Ecology 97: 10–17.
The indices characterizing plant colonization ability were published by Prach et al. (2017). The authors derived the index values for individual taxa from a database of 21 succession series (both primary and secondary succession) starting on the bare substrate. This database (Database of Successional Series, DaSS; Prach et al. 2014) contains 1013 taxa of vascular plants recorded in 2817 vegetation plots from the Czech Republic sampled in various habitats and successional stages of different age from 1 to 150 years. The following indices are defined:
Prach K., Tichý L., Vítovcová K. & Řehounková K. (2017) Participation of the Czech flora in succession at disturbed sites: quantifying species’ colonization ability. – Preslia 89: 87–100.
Chytrý M. & Rafajová M. (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. – Preslia 75: 1–15.
Prach K., Řehounková K., Lencová K., Jírová A., Konvalinková P., Mudrák O., Študent V., Vaněček Z., Tichý L., Petřík P., Šmilauer P. & Pyšek P. (2014) Vegetation succession in restoration of disturbed sites in Central Europe: the direction of succession and species richness across 19 seres. – Applied Vegetation Science 17: 193–200.
The floristic zones of the Earth in which the taxon occurs are defined according to Meusel et al. (1965, 1978) and Meusel & Jäger (1992). Data were taken from the BiolFlor database (Kühn & Klotz 2002).
Categories
Kühn I. & Klotz S. (2002) Angaben zu den Arealen. – In: Klotz S., Kühn I. & DurkaW. (eds), BIOLFLOR: eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland, Schriftenreihe für Vegetationskunde 38: 227–239.
Meusel H. & Jäger E. (1992) Vergleichende Chorologie der zentraleuropäischen Flora. Band III. – Gustav Fischer, Jena.
Meusel H., Jäger E. & Weinert E. (1965) Vergleichende Chorologie der zentraleuropäischen Flora. Band I. – Gustav Fischer, Jena.
Meusel H., Jäger E., Rauschert S. & Weinert E. (1978) Vergleichende Chorologie der zentraleuropäischen Flora. Band II. – Gustav Fischer, Jena.
The floristic region is reported as the continent or its part in which the taxon occurs according to the taxon range maps (Meusel et al. 1965, 1978, Meusel & Jäger 1992). The categories are not discrete, and some of the regions can be included within broader regions (e.g. Western Siberia – Siberia – Asia). From a set of overlapping categories, the one that best matches the taxon geographic range or its part is reported. Data were taken from the BiolFlor database (Kühn & Klotz 2002).
Kühn I. & Klotz S. (2002) Angaben zu den Arealen. – In: Klotz S., Kühn I. & DurkaW. (eds), BIOLFLOR: eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland, Schriftenreihe für Vegetationskunde 38: 227–239.
Meusel H. & Jäger E. (1992) Vergleichende Chorologie der zentraleuropäischen Flora. Band III. – Gustav Fischer, Jena.
Meusel H., Jäger E. & Weinert E. (1965) Vergleichende Chorologie der zentraleuropäischen Flora. Band I. – Gustav Fischer, Jena.
Meusel H., Jäger E., Rauschert S. & Weinert E. (1978) Vergleichende Chorologie der zentraleuropäischen Flora. Band II. – Gustav Fischer, Jena.
The lowest and the highest elevational vegetation belt in which the taxon commonly occurs in the Czech Republic. For some taxa, also extremes are shown, i.e. elevational belts in which the taxon rarely occurs outside its main elevational range. The submontane belt comprises merged supracolline and submontane belts, and the montane belt comprises merged montane and supramontane belts according to the classification of elevational vegetation belts used in the Flora of the Czech Republic (Skalický 1988). The data were taken from the Key to the Flora of the Czech Republic (Kaplan et al. 2019).
Kaplan Z., Danihelka J., Chrtek J. Jr., Kirschner J., Kubát K., Štěpánek J. & Štech M. (eds) (2019) Klíč ke květeně České republiky [Key to the flora of the Czech Republic]. Ed. 2. – Academia, Praha.
Skalický V. (1988) Regionálně fytogeografické členění [Regional phytogeographic division]. – In: Hejný S., Slavík B., Chrtek J., Tomšovic P. & Kovanda M. (eds), Květena České socialistické republiky [Flora of the Czech Socialist Republic] 1: 103–121, Academia, Praha.
The number of basic grid mapping cells (Central European Basic Area, CEBA) and the number of quadrants of the Central European flora mapping in that the taxon has been recorded within the territory of the Czech Republic are generated dynamically from the current occurrence records in the species distribution module of the Pladias Database. The basic grid cells measure 10 minutes in the west–east direction and 6 minutes in the south–north direction, which corresponds to approximately 12.0 × 11.1 km (133.2 km²) on the 50th parallel. The Czech Republic comprises 679 basic cells, including incomplete cells on the state borders. The quadrants are the basic grid cells divided into four. They measure 5 minutes in the west–east direction and 3 minutes in the south–north direction, which corresponds to approximately 6.0 × 5.55 km (33.3 km²) on the 50th parallel. Revised occurrence records marked as erroneous or uncertain are not counted.
Pladias. Database of the Czech flora and vegetation. – www.pladias.cz.
Measures of commonness in vegetation plots indicate taxon frequency in individual vegetation stands and the cover it attains. All these measures were quantified based on a set of vegetation plots representing all vegetation types of the Czech Republic, extracted from the Czech National Phytosociological Database (Chytrý & Rafajová 2003) in March 2013. These plots were classified to phytosociological associations using the expert system developed in the project Vegetation of the Czech Republic (Chytrý 2007–2013). The plots not assigned to any association were deleted, and a subset of plots of each association was selected based on a geographic stratification that reduced the unbalanced numbers of plots from different regions. The following measures of commonness were computed from the resulting set of 30,115 vegetation plots classified to 496 associations:
Chytrý M. (2016) Commonness in vegetation plots from the Czech Republic. – www.pladias.cz.
Chytrý M. (ed.) (2007–2013) Vegetace České republiky 1–4 [Vegetation of the Czech Republic 1–4]. – Academia, Praha.
Chytrý M. & Rafajová M. (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. – Preslia 75: 1–15.
The number of habitat types (habitats) in which the taxon occurs was counted based on the data from the Czech National Phytosociological Database (Chytrý & Rafajová 2003) and their expert revision and completion, especially for rare and taxonomically problematic taxa. This number is a measure of the taxon’s ecological range. The classification recognizes 88 basic habitats aggregated into 13 broader habitats that are defined by Sádlo et al. (2007: their Appendix 1). The number of habitats is defined in four ways:
Sádlo J., Chytrý M. & Pyšek P. (2007) Regional species pools of vascular plants in habitats of the Czech Republic. – Preslia 79: 303–321.
Chytrý M. & Rafajová M. (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. – Preslia 75: 1–15.
National Red List categories were taken from the 2017 edition of the Red List of Vascular Plants of the Czech Republic (Grulich 2017). These categories, introduced in the previous editions of the Czech Red List, are different from the IUCN Red List categories. The main category “A” includes extinct or missing taxa, while the main category “C” includes endangered, near threatened and data deficient taxa.
Grulich V. (2017) Červený seznam cévnatých rostlin ČR [The Red List of vascular plants of the Czech Republic]. – Příroda 35: 75–132.
International Red List categories defined by the IUCN were taken from the 2017 edition of the Red List of Vascular Plants of the Czech Republic (Grulich 2017). Taxon assignments to these categories follow the internationally accepted rules (IUCN 2012, 2014). To some extent, the definitions of these categories differ from the national categories used in the previous Czech Red Lists. The national Red List included only threatened or possibly threatened taxa, implying that the non-included taxa are not threatened. Therefore, the non-included taxa are classified here as LC(NA) – least concern (taxon is not on the Red List).
Grulich V. (2017) Červený seznam cévnatých rostlin ČR [The Red List of vascular plants of the Czech Republic]. – Příroda 35: 75–132.
IUCN (2012) Guidelines for application of IUCN Red List criteria at regional and national levels. Version 4.0. – IUCN, Gland.
IUCN (2014) Guidelines for using the IUCN Red List categories and criteria. Version 11. – IUCN, Gland.
Legal protection in the Czech Republic concerns the specifically protected species, i.e. rare taxa, threatened taxa and taxa significant from a cultural or scientific point of view that are listed in Annex II of the Decree of the Ministry of the Environment no. 395/1992. They comprise 487 taxa of vascular plants divided into three categories according to their vulnerability: critically threatened, endangered and vulnerable.
Decree no. 395/1992 of the Ministry of the Environment of the Czech Republic.