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.
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.
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.
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.
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.
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.
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.
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.
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.
Invasion status is a classification of alien taxa into three categories reflecting their position in the invasion process. Alien taxa that only occasionally reproduce in the wild in the Czech Republic, do not form self-replacing populations, and rely on repeated introductions for their persistence are termed casual. Naturalized taxa are alien plants that reproduce in the wild and sustain populations over many life cycles without direct intervention by humans (or despite human intervention). Invasive plants are naturalized plants that produce reproductive offspring, often in large numbers, at considerable distances from parent plants and thus have the potential to spread over an extensive area (Richardson et al. 2000, 2011). This classification does not apply to native taxa, which are reported as separate categories. The data were taken from 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. & Carlton J. T. (2011) A compendium of essential concepts and terminology in biological invasions. – In: Richardson D. M. (ed.), Fifty years of invasion ecology: the legacy of Charles Elton, p. 409–420, Blackwell Publishing, Oxford.
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.
This information is given for alien taxa only. These taxa are classified according to their geographic origin (native range) at the level of continents; those with a native range encompassing more than one continent are assigned to two or more categories. Origin in Europe refers to the non-Mediterranean parts of this continent other than the Czech Republic. The Mediterranean region comprises parts of southern Europe, northern Africa and western Asia from Turkey and Israel to Afghanistan, which are characterized by the Mediterranean-type climate and the occurrence of sclerophyllous evergreen vegetation. Conversely, records of origin in Africa, Asia and Europe do not relate to the Mediterranean part of these continents. Hybrids and species that originated through recent hybridization are listed as a separate category. Anecophytes are taxa for which native range is unknown or highly uncertain. The data were taken from 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.
The year of the first reported occurrence in the wild in the Czech Republic is given for neophytes. Data were extracted from the third edition of the Catalogue of alien plants of the Czech Republic (Pyšek et al. 2022); however, for many species, this information is not available.
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.
Period of introduction is divided into categories from the Neolithic to the present time. The assessment is based on archaeobotanical and also floristic data where applicable. For taxa without precise dating, only merged categories of longer time periods are provided.
Data were extracted from the third edition of the Catalogue of alien plants of the Czech Republic (Pyšek et al. 2022).
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.
This information is available for alien taxa only. Taxa are classified according to the mode of introduction, with the distinction made between an intentional (deliberate) and unintentional (accidental) pathway (Hulme et al. 2008). A taxon can be assigned to more than one introduction pathway. The pathway classification and data for individual taxa follow the third edition of the Catalogue of alien plants of the Czech Republic (Pyšek et al. 2022).
The classification includes the following introduction pathways:
Intentional introduction: 1 ornamental – cultivation for ornamental purposes in a garden or public green space; 2 collection – cultivation for scientific or collection purposes in a garden or park; 3 nature – ornamental planting in natural habitats, including deceptive enrichment of regional flora or inclusion of alien plants in seed mixtures sold as “wild meadows”; 4 crops – cultivation of crops such as fruits, vegetables, cereals, spices; 5 forest – forestry and wildlife management, e.g. planting trees in forestry plantations or cultivation of crops in forests as a fodder for game; 6 landscaping, e.g. afforestation, land reclamation, roadside tree belts, tree alleys; 7 other – cultivation for other purposes, e.g. forage crops, plants for basketry, beekeeping and pharmacy.
Unintentional introduction: 8 agriculture – introduction associated with agriculture: weeds in fields (e.g. cereals, flax), vegetable beds and vineyards; 9 plantation – introduction associated with fodder crop cultivation, sowing meadows and planting forest trees; 10 horticulture – introduction associated with the horticultural sale of ornamental plants, i.e. weeds of ornamental gardens; 11 traffic – dispersal by vehicles, trains, shipping, urban tourism; 12 industry – dispersal through mining or production/distribution of commodities, e.g. ores, wool; 13 humans – other unintended vectors associated with anthropogenic habitats; 14 nature – other unintended vectors associated with natural habitats, e.g. dispersal by wind, water, animals.
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.
Hulme P. E., Bacher S., Kenis M., Klotz S., Kühn I., Minchin D., Nentwig W., Olenin S., Panov V., Pergl J., Pyšek P., Roques A., Sol D., Solarz W. & Vilà M. (2008) Grasping at the routes of biological invasions: a framework for integrating pathways into policy. – Journal of Applied Ecology 45: 403–414.
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.