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The results of a taxonomic study ofCardamine flexuosa
With.,C. glauca
Spreng. ex DC.,C. graeca L.,C. hirsuta L.,C. impatients L.,C. parviflora L.,C. resedifolia L., andC. trifolia L. from the Carpathians and Pannonia are presented. Full synonymy, lectotypifications, descriptions, chromosome numbers,
taxonomic history, notes on variation, distributi...
Citations
... The chromosome count in the Brassicaceae is an efficient tool used to set up the boundaries of the relations among taxa within genera and they are of great significance when reviewing systematics and evolution in this family (Karaismailoğlu 2018). Some chromosome numbers and morphological investigations in the genus Cardamine from various parts of the world have been reported (Hussein 1948, Berg 1966, Thurling 1968, Marhold 1994, 1995, Kučera et al. 2005, Perny et al. 2005, Warwick and Al-Shehbaz 2006. However, the taxonomic usability of chromosome number in the genus has been ignored in Turkey, until now. ...
... Several attempts have been made to split this polymorphic species into more entities. Some authors recognized Cardamine nemorosa and Cardamine udicola in lowland to montane areas (Urbanska-Worytkiewicz and Landolt, 1974), and diploid populations from the foothills of the Eastern Carpathians were informally treated as a morphotype 'ucranica' (Marhold, 1994b(Marhold, , 1995. In addition, high-elevation Alpine and Eastern Carpathian populations were attributed to C. rivularis by Lövkvist (1956) and Urbanska-Worytkiewicz and Landolt (1974; see Table 1). ...
... Cardamine matthioli is a diploid (2n = 16) distributed in Central Europe and extending to Southeastern Europe (Marhold, 1994b(Marhold, , 2000. Aneuploids with supernumerary chromosomes ranging from 17 to 22 have been observed sporadically (Kučera et al., 2005). ...
... Aneuploids with supernumerary chromosomes ranging from 17 to 22 have been observed sporadically (Kučera et al., 2005). The species grows mainly in lowlands and up to the montane belt, and only exceptionally reaches elevations above Marhold et al. (2018) and present study Ploidy level (Kučera et al., 2005) Distribution range Lövkvist, 1956Urbanska-Worytkiewicz and Landolt, 1974Landolt, 1984Marhold, 1994a Franzke 1,000-1,500 m (Marhold, 1994b;Ančev, 2006). In total, 223 plants from 28 populations were collected. ...
Recurrent polyploid formation and weak reproductive barriers between independent polyploid lineages generate intricate species complexes with high diversity and reticulate evolutionary history. Uncovering the evolutionary processes that formed their present-day cytotypic and genetic structure is a challenging task. We studied the species complex of Cardamine pratensis, composed of diploid endemics in the European Mediterranean and diploid-polyploid lineages more widely distributed across Europe, focusing on the poorly understood variation in Central Europe. To elucidate the evolution of Central European populations we analyzed ploidy level and genome size variation, genetic patterns inferred from microsatellite markers and target enrichment of low-copy nuclear genes (Hyb-Seq), and environmental niche differentiation. We observed almost continuous variation in chromosome numbers and genome size in C. pratensis s.str., which is caused by the co-occurrence of euploid and dysploid cytotypes, along with aneuploids, and is likely accompanied by inter-cytotype mating. We inferred that the polyploid cytotypes of C. pratensis s.str. are both of single and multiple, spatially and temporally recurrent origins. The tetraploid Cardamine majovskyi evolved at least twice in different regions by autopolyploidy from diploid Cardamine matthioli. The extensive genome size and genetic variation of Cardamine rivularis reflects differentiation induced by the geographic isolation of disjunct populations, establishment of triploids of different origins, and hybridization with sympatric C. matthioli. Geographically structured genetic lineages identified in the species under study, which are also ecologically divergent, are interpreted as descendants from different source populations in multiple glacial refugia. The postglacial range expansion was accompanied by substantial genetic admixture between the lineages of C. pratensis s.str., which is reflected by diffuse borders in their contact zones. In conclusion, we identified an interplay of diverse processes that have driven the evolution of the species studied, including allopatric and ecological divergence, hybridization, multiple polyploid origins, and genetic reshuffling caused by Pleistocene climate-induced range dynamics.
... Our study sites were in the areas inhabited by only diploids. Cardamine hirsuta L. is a diploid (2n = 2x = 16) native to Europe (Marhold, 1995;Lihová et al., 2006;Grime et al., 2007). It is a winter annual and its seeds cannot tolerate submergence (Yatsu et al., 2003). ...
Polyploidization is pervasive in plants, but little is known about the niche divergence of wild allopolyploids (species that harbor polyploid genomes originating from different diploid species) relative to their diploid progenitor species and the gene expression patterns that may underlie such ecological divergence. We conducted a fine‐scale empirical study on habitat and gene expression of an allopolyploid and its diploid progenitors.
We quantified soil properties and light availability of habitats of an allotetraploid Cardamine flexuosa and its diploid progenitors Cardamine amara and Cardamine hirsuta in two seasons. We analyzed expression patterns of genes and homeologs (homeologous gene copies in allopolyploids) using RNA sequencing.
We detected niche divergence between the allopolyploid and its diploid progenitors along water availability gradient at a fine scale: the diploids in opposite extremes and the allopolyploid in a broader range between diploids, with limited overlap with diploids at both ends. Most of the genes whose homeolog expression ratio changed among habitats in C. flexuosa varied spatially and temporally.
These findings provide empirical evidence for niche divergence between an allopolyploid and its diploid progenitor species at a fine scale and suggest that divergent expression patterns of homeologs in an allopolyploid may underlie its persistence in diverse habitats.
... Tapasztalataink szerint az utóbbi fajhoz képest ritkábban fordul elő a kertészetekben: 13 helyen találtuk. Míg korábban ismeretlen volt az Alföldön (MARHOLD 1995), újabban előkerült néhány előfordulása [3]. Első tiszántúli (Szentes) felfedezése kapcsán JAKAB (2005) vetette föl, hogy tőzeges virágfölddel hurcolhatták be. ...
During the study of the weed flora of garden centers in Hungary (among 2017–2020), remarkable populations of Cardamine occulta Hornem., a new alien for the Hungarian flora were found. C. occulta was present altogether in 51 of the 53 visited sites. Dominantly the regularly irrigated and continuously moist microhabitats (pots, containers, muddy surfaces of geotextile-covered beds etc.) were colonized. During the revision of our recently collected specimens, deposited in JPU and DE herbaria as Cardamine hirsuta L., further individuals proved to identical with this till overlooked species. One of them (27.08.2004., Heves county: Eger [8088.3; 8188.1], coll. by A. Schmotzer, deposited in DE collection) proved to the third documented occurrence in Europe, comparing to the accessed literature data. Eclipta prostrata (L.) L. and Urtica membranacea Poir. are also new aliens for the Hungarian flora. Several introduced individuals of these taxa were found in containers of imported thermophilous woody ornamentals at 4 and 2 sites, respectively. Tens of individuals of Eclipta prostrata were also found in a sapling-bed at another site. New populations of scarce or rare Euphorbia prostrata Aiton, E. serpens Kunth and Veronica peregrina L. were also documented.
... In Switzerland, the populations of C. amara in lowland areas (< 1,000 m) are reported to consist of diploids (2n = 2x = 16), and autotetraploid populations of the subsp. austriaca have been observed at higher altitudes in the inner Alpine valleys (Marhold 1995(Marhold , 1999Marhold et al. 2002). Cardamine amara propagates clonally and sexually (Tedder et al. 2015b). ...
... Cardamine hirsuta L. (hairy bittercress) is a diploid (2n = 2x = 16) annual herb native to Europe (Grime et al. 2007;Lihová et al. 2006b;Marhold 1995;Yatsu et al. 2003). It grows 5-30 cm in height and forms a rosette (Lauber et al. 2012). ...
Allopolyploids possess complete sets of genomes derived from different parental species and exhibit a range of variation in various traits. Reproductive traits may play a key role in the reproductive isolation between allopolyploids and their parental species, thus affecting the thriving of allopolyploids. However, empirical data, especially in natural habitats, comparing reproductive trait variation between allopolyploids and their parental species remain rare. Here, we documented the flowering phenology and floral morphology of the allopolyploid wild plant Cardamine flexuosa and its diploid parents C. amara and C. hirsuta in their native range in Switzerland. The flowering of C. flexuosa started at an intermediate time compared with those of the parents and the flowering period of C. flexuosa overlapped with those of the parents. Cardamine flexuosa resembled C. hirsuta in the size of flowers and petals and the length/width ratio of petals, while it resembled C. amara in the length/width ratio of flowers. These results provide empirical evidence of the trait-dependent variation of allopolyploid phenotypes in natural habitats at the local scale. They also suggest that the variation in some reproductive traits in C. flexuosa is associated with self-fertilization. Therefore, it is helpful to consider the mating system in furthering the understanding of the processes that may have shaped trait variation in polyploids in nature.
... The generalist niche and transcriptomic plasticity of an allopolyploid we documented here in C. flexuosa can be common to other allopolyploid species. For example, the genus but also in other allopolyploid species, such as cotton, coffee, and Arabidopsis (5,9,13,61,62). Both in the present study (Table S5) and previous studies on Brassicaceae in the laboratory (10,12), only a small portion (<4%) of all genes exhibited the homeolog . CC-BY-NC-ND 4.0 International license It is made available under a (which was not peer-reviewed) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. ...
Polyploidization, or whole genome duplication, is one of the major mechanisms of plant speciation. Allopolyploids (species that harbor polyploid genomes originating from hybridization of different diploid species) have been hypothesized to occupy a niche with intermediate, broader, or fluctuating environmental conditions compared with parental diploids. It remains unclear whether empirical data support this hypothesis and whether specialization of expression patterns of the homeologs (paralogous gene copies resulting from allopolyploidization) relates to habitat environments. Here, we studied the ecology and transcriptomics of a wild allopolyploid Cardamine flexuosa and its diploid parents C. hirsuta and C. amara at a fine geographical scale in their native area in Switzerland. We found that the diploid parents favored opposite extremes in terms of soil moisture, soil carbon-to-nitrogen ratios, and light availability. The habitat of the allopolyploid C. flexuosa was broader compared with those of its parental species and overlapped with those of the parents, but not at its extremes. In C. flexuosa , the genes related to water availability were overrepresented among those at both the expression level and the expression ratio of homeolog pairs, which varied among habitat environments. These findings provide empirical evidence for niche differentiation between an allopolyploid and its diploid parents at a fine scale, where both ecological and transcriptomic data indicated water availability to be the key environmental factor for niche differentiation.
Significance statement
Polyploidization, or whole genome duplication, is common in plants and may contribute to their ecological diversification. However, little is known about the niche differentiation of wild allopolyploids relative to their diploid parents and the gene expression patterns that may underlie such ecological divergence. We detected niche differentiation between the allopolyploid Cardamine flexuosa and its diploid parents C. amara and C. hirsuta along water availability gradient at a fine scale. The ecological differentiation was mirrored by the dynamic control of water availability-related gene expression patterns according to habitat environments. Thus, both ecological and transcriptomic data revealed niche differentiation between an allopolyploid species and its diploid parents.
... Nevertheless, it is worth commenting here on the differences between C. hirsuta and the species studied here. Whereas C. flexuosa, C. kokaiensis, C. occulta and C. scutata always have six stamens, plants of C. hirsuta predominantly have four (Marhold, 1995; but see Matsuhashi, Sakai & Kudoh, 2012). Other characters that characterize C. hirsuta are a completely glabrous and straight stem and a compact rosette of basal leaves. ...
Cardamine occulta, recently reported also as 'Asian C. flexuosa', is originally a native East Asian weed of paddy fields and other open habitats. It has been recorded throughout the world and is now also spreading widely in Europe. However, how this species differs morphologically from European C. flexuosa and its closest relatives in Asia (C. kokaiensis and C. scutata) is not fully known, particularly because it became widely recognized as a new taxon only recently. We used chromosome counting and flow cytometry to determine ploidies and genome size variation in these four species and morphometric analyses to ascertain their morphological differentiation. A uniformly octoploid level (2n = 64) is confirmed here for C. occulta and a tetraploid level for C. flexuosa and C. scutata (2n = 32). Here we formally describe C. kokaiensis, which is restricted to East Asia (Japan, Honshu; eastern China, Zhejiang Province; perhaps also the Russian Far East) and determine it to be a tetraploid. Considerable differences in monoploid genome size were revealed between these taxa, suggesting their mostly allopolyploid origins. Morphological differences between the species are demonstrated, and detailed morphological descriptions and an identification key are provided.
... Such habitat conditions are particularly suitable because the species prefers periodically fl ooded and open lowland areas (ZLINSKÁ 1990). The small-fl owered bittercress is already known from certain communities of alliances such as Nanocyperion Koch 1926, Phalaridion arundiaceae Kopecky 1961or Phragmition communis Koch 1926(ZLINSKÁ 1992, MARHOLD 1995. ...
Cardamine parviflora L. was discovered in April 2014 during the study of vascular flora and habitats in the area of Slatina (Slavonia region). It was found in a flooded forest of narrow-leaved ash, in the vicinity of the villages Medinci and Novi Senkovac. The species here grows in wet soil but partially submerged populations were also observed. It presents a new species of the Croatian flora and expands the floristic inventory of Slatina and its surroundings. Other valuable taxa have also been recorded in the area, such as Carex riparia Curtis and Ophioglossum vulgatum L.
... The species collected for the first time by Sprengel in 1819 ("In aspero monte prope Reggio, in Callabria") (Marhold, 1995). The first official description of the species was made in 1821 by Augustin Pyramus de Candolle in Regni Vegetabilis Systema Naturale (Candole, 1821) ( Fig. 1). ...
... Bibliographical data about Cardamine glauca from romanian botanical literature is presented in Table 1. It is worth mentioning that location "Căpăţânii Mountains, between Novaci and Şugag, on the mountain ridge" (Marhold, 1995;Oprea, 2005) is not plausible, both localities Novaci and Şugag are not situated near Căpăţânii Mountains. Novaci locality is situated near Parâng Mountains and Şugag near Şureanu Mountains. ...
Cardamine glauca Spreng. ex DC. is a Critically Endangered, Balkanic-Appenninic-Dacian floristic element, distributed in Carpathians only in Parâng and Făgăraş Mountains. Based on field studies, analyses of herbarium material and literature data, the authors managed to record the occurrence of Cardamine glauca in Carpathians and determined the threatened status according to criteria and categories of IUCN. The authors have discovered two new localities of this important European endemic species (in Făgăraş Mountains).
... Algunas de sus especies muestran una elevada complejidad taxonómica, como es el caso de C. flexuosa With., sumamente polimorfa (MARHOLD, 1995;RICO, 1996) y muy próxima a C. hirsuta, congénere con el que en ocasiones es difícil distinguir, y que también muestra una gran variabilidad. Además, estas dos especies pueden hibridar, lo que complica la determinación de algunos ejemplares cuando ambas conviven (RICO, 1996;ELLIS & JONES, 1969). ...
... Trabajos taxonómicos posteriores no reconocen esta división, incluyendo el material procedente de estas dos metapoblaciones bajo el mismo nombre. No obstante, ambas metapoblaciones muestran una extraordinaria plasticidad fenotípica (KUDOH & al., 1995;MARHOLD, 1995;RI-CO, 1996;WU & al., 2001) y a pesar de ello, se reconoce un único taxon de amplia variabilidad morfológica. Según MARHOLD (1995:400), la variación de las subpoblaciones europeas no merece reconocimiento taxonómico, y con respecto a las subpoblaciones orientales sugiere que probablemente pertenezcan a otro taxón próximo. ...
Taxonomic notes on Cardamine flexuosa With. (Cruciferae)
complex and its presence in the Valencian Community. First record of Cardamine
flexuosa With. s. str. from the Valencian Community and first record of Cardamine
flexuosa subsp. debilis O. E. Schulz from the Province of Valencia, both
placed at Eastern Spain, are reported. Some nomenclatural, taxonomic, chorological
and ecological aspects reporting both taxa are reviewed, as well as those related to
Cardamine hirsuta L., a morphologically very similar species. A dichotomous key
to identify and separate these three taxa is proposed. The introduction way of the alien
species C. flexuosa s.l. is suggested and an evaluation of the status as a possible
potential invasive species is made. Key words: Cardamine flexuosa subsp. debilis,
taxonomy, alien species, Valencian Community, Cruciferae, Spain.
