The genus Rubus is among the most taxonomically complex and species-rich groups of vascular plants in Europe, owing to extensive morphological variation, polyploidy, facultative apomixis and frequent hybridisation. An updated checklist of the genus Rubus in Poland, presented here, comprises 113 taxa, including 106 native species and seven established alien taxa; 12 species occur exclusively in Poland and are treated as regional endemics. An illegitimate later homonym, Rubus magurensis , is replaced here with the new name, Rubus subbeskidensis . Using 88,934 verified records from the ATPOL database, spatial patterns of species richness are analysed and seven regional micro-hotspots of Rubus diversity in Poland are identified. The highest concentrations of species are located in south-western Poland, particularly in the Kotlina Kłodzka Basin, the Moravian Gate and the forelands of the northern Carpathians. Phytogeographical analyses of the distribution patterns of all brambles occurring in Poland allow the recognition of four distinct geographical elements, comprising species reaching their northern, north-eastern, western and southern distribution limits in Poland. The distribution patterns of selected species groups depend mainly on specific bioclimatic factors determining the occurrence of individual species.
Rubus ser. Glandulosi is one of the most taxonomically intricate taxa of the genus, which is by itself considered a taxonomical nightmare. Around 46 species are currently recognised in the group, but most of its morphological diversity could not be classified at the species level until recently due to unclear reproductive patterns, high genotypic diversity, low morphological differentiation among genotypes, and high phenotypic plasticity. This contribution utilises the recent findings in biosystematics of this taxon and aims at a natural classification reflecting its evolution. First, we explicitly define two different species concepts for sexual and apomictic populations. Second, we utilise microsatellite genotyping to identify widely distributed apomictic genotypes, some of which can be linked with currently recognised species. To this end, we genotyped 31 nomenclatural types and 14 “topotype” specimens or specimens of the original material. For the sexual species and apomictic microspecies whose genotypic homogeneity, considerable distribution, and identity with the nomenclatural type were studied and confirmed by genotyping, we provide morphological diagnostic characters and photographic documentation. Epitypes for the names R. flaccidifolius , R. guentheri , and R. lusaticus are designated. Three morphologically identifiable and molecularly characterised genotypes are described as new species – R. coronae‐aureae , R. divulgatus , and R. piceeticola . A complete annotated checklist of currently recognised species and a simplified determination key are provided. Several older names are further proposed for exclusion due to their probably low taxonomic value.
Hybridization and polyploidization are significant evolutionary mechanisms. Allopolyploidy, the formation of polyploids through hybridization between distinct species followed by whole-genome duplication, is common mainly in plants, where it facilitates rapid speciation and ecological diversification. Hybridization between diploid species often results in F1 hybrids with reduced fertility due to irregular meiosis. However, subsequent genome duplication can restore fertility by providing homologous chromosome pairs. Understanding the initial hybridization between diploids is therefore crucial for elucidating the early stages of allopolyploid formation. Ranunculus sect. Batrachium is known for the frequent occurrence of hybrids and polyploids, many of which are allopolyploid. Surprisingly, no unequivocal natural hybrid between diploid species has been discovered until recently in spite of the large body of evidence on interspecific hybridization in this group. This study provides evidence for hybridization at the diploid level, based on an integrated morphological, cytological and molecular approach. A new interspecific hybrid R. circinatus x R. rionii was detected in two flooded sand pits in the Czech Republic and described as R. xlimnophilus. Maternal inheritance patterns indicated independent, reciprocal origin of the hybrids at each locality. The ecological and historical context of the hybrid's emergence is discussed, including colonization of a newly formed habitat that allowed coexistence of the two parental species, their hybridization and the subsequent vegetative persistence of the hybrid despite its sterility. Similar conditions may have prevailed during the early postglacial period, facilitating the formation of allopolyploids, which today constitute the majority of species diversity in this group. A review and categorization of the biotypes that constitute the complex diversity in Ranunculus sect. Batrachium is also provided.
AbstractRubusser.Glandulosirepresents a unique model of geographic parthenogenesis on a homoploid (4x) level. We employed double digest restriction-site associated DNA sequencing (ddRADseq) in 587 individuals of differentRubustaxa and modelling of suitable climate to characterize evolutionary and phylogeographic patterns and shed light on the geographic differentiation of apomicts and sexuals. Six ancestral species were identified that contributed to the contemporary gene pool ofR.ser.Glandulosi. While sexuals were introgressed fromR. dolichocarpusandR. moschusin West Asia and fromR. ulmifoliusagg.,R. canescensandR. incanescensin Europe, apomicts were characterized by alleles ofR.subsect.Rubus. Gene flow between sexuals and apomicts was also detected, as well as occasional hybridization with other taxa. We hypothesize that sexuals survived the last glacial period in several large southern refugia, whereas apomicts were mostly restricted to southern France from whence they quickly recolonized Central and Western Europe. The secondary contact of sexuals and apomicts was probably the principal factor that established geographic parthenogenesis inR.ser.Glandulosi. Sexual populations are not impoverished in genetic diversity along their borderline with apomicts and maladaptive population genetic processes likely did not shape the geographic patterns.HighlightsGeographic parthenogenesis inRubusser.Glandulosiis caused by secondary contactReticulate evolution in Eurasian brambles is more extensive than expectedAt least six species contributed to the evolution ofRubusser.GlandulosiExtinct ancestor’s gene pool is associated with apomixisAbstract FigureGraphical abstract
Background and Aims Rubus ser. Glandulosi provides a unique model of geographical parthenogenesis on a homoploid (2n = 4x) level. We aim to characterize evolutionary and phylogeographical patterns in this taxon and shed light on the geographical differentiation of apomicts and sexuals. Ultimately, we aim to evaluate the importance of phylogeography in the formation of geographical parthenogenesis.Methods Rubus ser. Glandulosi was sampled across its Eurasian range together with other co-occurring Rubus taxa (587 individuals in total). Double-digest restriction site-associated DNA sequencing (ddRADseq) and modelling of suitable climate were used for evolutionary inferences.Key Results Six ancestral species were identified that contributed to the contemporary gene pool of R. ser. Glandulosi. Sexuals were introgressed from Rubus dolichocarpus and Rubus moschus in West Asia and from Rubus ulmifolius agg., Rubus canescens and Rubus incanescens in Europe, whereas apomicts were characterized by alleles of Rubus subsect. Rubus. Gene flow between sexuals and apomicts was also detected, as was occasional hybridization with other taxa.Conclusions We hypothesize that sexuals survived the last glacial period in several large southern refugia, whereas apomicts were mostly restricted to southern France, whence they quickly recolonized Central and Western Europe. The secondary contact of sexuals and apomicts was probably the principal factor that established geographical parthenogenesis in R. ser. Glandulosi. Sexual populations are not impoverished in genetic diversity along their borderline with apomicts, and maladaptive population genetic processes probably did not shape the geographical patterns.
Asexual organisms often differ in their geographic distributions from their sexual relatives. This phenomenon, termed geographic parthenogenesis, has long been known, but the underlying factors behind its diverse patterns have been under dispute. Particularly problematic is an association between asexuality and polyploidy in most taxa. Here, we present a new system of geographic parthenogenesis on the tetraploid level, promising new insights into this complex topic. We used flow cytometric seed screen and microsatellite genotyping to characterise the patterns of distribution of sexuals and apomicts and genotypic distributions in Rubus ser. Glandulosi across its range. Ecological modelling and local-scale vegetation and soil analyses were used to test for niche differentiation between the reproductive groups. Apomicts were detected only in North-western Europe, sexuals in the rest of the range in Europe and West Asia, with a sharp borderline stretched across Central Europe. Despite that, we found no significant differences in ecological niches. Genotypic richness distributions suggested independence of the reproductive groups and a secondary contact. We argue that unless a niche differentiation (resulting from polyploidy and/or hybridity) evolves, the main factors behind the patterns of geographic parthenogenesis in plants are phylogeographic history and neutral microevolutionary processes, such as clonal turnover.
Effective protection of endangered species is often limited by taxonomic discrepancies across state borders. This is also the case of the Dactylorhiza maculata agg. in Central Europe, where one to three species and several infraspecific taxa are recognized in various countries. Based on an extensive analysis of morphological variation, ploidy levels, environmental traits and habitats of 64 populations in Central Europe and adjacent regions, we aimed to propose a unified taxonomic concept applicable throughout the study area. Multivariate analysis of morphological traits revealed continuous variation at the individual level and only minor differences between particular clusters of populations. Four DNA-ploidy levels were detected using flow cytometry. Diploids (2 n = 40) and tetraploids (2 n = 80) were the most abundant and usually formed single-cytotype populations whereas DNA-triploids and DNA-hexaploids occurred only sporadically as minority cytotypes. The inferred patterns of morphological and ploidy variation were not congruent with traditional taxonomic treatment regarding diploid D. fuchsii and tetraploid D. maculata as two species with several infraspecific taxa. Instead, all taxa analysed in the current study are best treated at the subspecies level within D. maculata s. lat. due to somewhat continuous morphological variation between morphotypes. A total of eight D. maculata subspecies may be recognized in Central Europe, of which one is newly described here as D. maculata subsp. arcana , subsp. nov. Some nomenclatural riddles have been resolved, and the threat status of the recognized taxa is discussed.
Semi-dry grasslands are among the most species-rich plant communities in the world, harbouring many specialised and threatened species. Most of these grasslands were traditionally maintained by grazing and hay-making. After traditional management ended, protected areas were established and conservation management was introduced to protect the most valuable grassland sites. However, recent changes in land use, eutrophication and climate warming are negatively impacting the biodiversity of these grasslands. In 2022, we resurveyed historical vegetation plots in the Central Moravian Carpathians (Czech Republic), first sampled in the 1980s, to test whether the plant species composition and richness of semi-dry grasslands are changing over time and, if so, whether the decline in habitat quality and plant diversity is absent or less severe in protected areas. We found significant changes in species composition. Species richness and the proportion of habitat specialists and Red-List species decreased, whereas competitively stronger species with higher moisture and nutrient requirements increased. These trends were more pronounced outside the protected areas but also occurred within protected areas. The main factor behind these changes appears to be the cessation of traditional management and natural succession supported by eutrophication.
(2963) Rubus pseudoidaeus F.W. Schmidt in Neue Abh. Königl. Böhm. Ges. Wiss. 1: 37. 1790, nom. rej. prop. Lectotypus (vide Van de Beek in Gorteria 39: 52. 2017): “Rubus schmidtianus Presl / R. pseudoidaeus Schmidt. fl. boh. / In sepibus ad Moldavam superiorem” (PRC). Rubus pseudoidaeus F.W. Schmidt (in Neue Abh. Königl. Böhm. Ges. Wiss. 1: 37. 1790) is a very early published name of a European Rubus species. Van de Beek (in Gorteria 39: 52. 2017) argued that it is a heterotypic earlier synonym of Rubus sulcatus Vest (in Steiermärk. Z. 3: 162. 1821), a well-known species with a wide distribution. The name Rubus pseudoidaeus F.W. Schmidt has never been used in later literature, while R. sulcatus was accepted by Trattinnick (Rosac. Monogr. 3: 42. 1823), and has subsequently been used by all experts in the genus Rubus, especially after Focke (Syn. Rub. Germ.: 119. 1877) acknowledged its identity. Rubus sulcatus is not just one of the many Rubus taxa with a limited local or regional distribution. It is one of the few European taxa found in all temperate regions of Europe, including the British Islands (Kurtto & al., Atlas Fl. Eur.: 52. 2010). These obviously old taxa are not only of interest for their distribution but also for understanding the propagation processes within Rubus. They have very similar taxa along the type of which the origin is unclear. Much research on this topic is still ahead, and stability in nomenclature is even more critical in such a situation. The epithet ‘pseudoidaeus’ has been in general use for a totally different taxon: the hybrid of Rubus caesius L. and R. idaeus L., first at the rank of variety (Weihe in Boenninghausen, Prodr. Fl. Monast. Westphal.: 151. 1824), and soon as a species (Schlechtendahl in Linnaea 8: 315. 1833) (usually botanists, e.g., Weber, Rubi Westphalici: 426. 1985, refer to Lejeune [Rev. Fl. Spa: 102. 1825] as the author, but that publication does only refer to an unpublished [“ined.”] name of a specimen). Because changing the well-known name Rubus sulcatus to R. pseudoidaeus, which has never been in use for this taxon since its publication in 1790, would cause much confusion, Van de Beek (in Gorteria 39: 52. 2017) suggested submitting a proposal for the conservation of the name R. sulcatus. However, the taxa of Rubus ser. Rubus, to which R. sulcatus belongs, have relatively few distinctive characters, and such similar – but not identical – taxa as indicated above, were found in the precise region of the type locality of R. pseudoidaeus F.W. Schmidt, such as R. altissimus Fritsch (in Hayek, Sched. Fl. Stiriac. 5–6: 11. 1905) and R. barrandienicus Holub & Palek (in Folia Geobot. Phytotax. 26: 332. 1991), so that, eventually, it might turn out that R. pseudoidaeus is not identical with R. sulcatus. Therefore, to avoid any imminent or future confusion, we propose rejecting outright the name Rubus pseudoidaeus F.W. Schmidt. AvdB, https://orcid.org/0000-0001-6728-9572 DD, https://orcid.org/0009-0005-3976-1020 HDV, https://orcid.org/0000-0003-4011-8225 TG, https://orcid.org/0000-0001-7692-8348 EvdH, https://orcid.org/0009-0002-6578-1509 RH, https://orcid.org/0000-0001-9127-4549 MH, https://orcid.org/0000-0002-3880-9417 GK, https://orcid.org/0000-0002-8439-2616 PK, https://orcid.org/0000-0001-6104-4267 GM-H, https://orcid.org/0009-0001-6093-8768 KM, https://orcid.org/0009-0005-5113-461X DPM, https://orcid.org/0009-0008-2308-0783 IdR, https://orcid.org/0000-0002-6008-6956 UR, https://orcid.org/0000-0001-7653-8489 BT, https://orcid.org/0000-0003-4596-2476
Gagea bohemica s.l. is a morphologically and karyologically highly variable group with many morphologically similar “narrow” taxa currently considered as a single variable species. It is predominantly distributed in Mediterranean and warmer parts of temperate belt of Europe. The large-scale data on its cytogeography and population cytotype structure which could provide a basis for taxonomy are lacking, only scattered data on ploidy have been published from various parts of its range. In this study, we sampled 106 populations in broader Central Europe, the northeastern Balkan Peninsula and the northwestern Black Sea coast in order to analyze their ploidy level, genome size and pollen stainability. Two cytotypes, i.e., tetraploid (2 n = 48) and pentaploid (2 n = 60), were found in the study area using chromosome counting and flow cytometry, both in pure and mixed-ploidy populations. Pure pentaploid populations are mainly distributed in Austria, Czechia, northwestern Hungary and Slovakia while tetraploid cytotype in pure and mixed-ploidy populations forming two lineages which are concentrated into two disjunct geographical areas: a western lineage in Germany and Switzerland and an eastern one in Bulgaria, southeastern Hungary, northern Greece, Romania, Serbia and Ukraine. The two lineages differ in their genome size regardless of their ploidy, indicating their independent origin. Analysis of pollen stainability using a modified Alexander stain revealed an unusual pattern with tetraploids having a lower pollen stainability (mean 44.29%) than pentaploids (mean 70.70%), but the western and eastern populations differed again from each other.
According to currently available data, 159 species of genus Taraxacum F. H. Wigg. were reported for North-West Russia (i. e. within the city of Saint-Petersburg, Leningrad Region, Pskov Region, and Novgorod Region), but among them the specimens of only 126 species were identified with reliability. Five Taraxacum species are reported in this article as new for North-West Russia, and for Russia as a whole: T. acervatulum Rail., T. hepaticum Rail., T. macranthoides G. E. Haglund, T. sertatum Kirschner, H. Øllg. et Štěpánek, T. subxanthostigma M. P. Christ. ex H. Øllg. All of them belong to the section Taraxacum. Newly recorded species are illustrated and the list of their herbarium specimens is provided. A brief summary of the taraxacological studies undertaken earlier in North-West Russia is presented as well.
Rubus violaceifrons, a new species from central and north-eastern Bohemia (Czech Republic) is described and illustrated, and its distribution is characterized in detail. We classify it as a member of the series Pallidi of the subgenus Rubus, section Rubus and subsection Hiemales, although its weak specimens partly resemble R. guentheri from ser. Glandulosi. Like all species of R. ser. Pallidi studied so far, R. violaceifrons is tetraploid, as determined using flow cytometry. A detailed morphological comparison of the new bramble species with similar taxa is presented. A determination key including all taxa of the R. ser. Pallidi occurring in the Czech Republic is provided.
Rubus ser. Subthyrsoidei is a particularly intricate group within the sect. Corylifolii due to its phenologically and ecologically driven plasticity and variability. In the recent study we investigated the narrower-leaved biotypes of the series in south-eastern central Europe that represent a distinct entity often called “Rubus gothicus group”. We recognized four stable biotypes (apomictic species) occurring in the area: (1) the north-central European R. gothicus s. str., which was rarely found in northern Bohemia; (2) a biotype from south-eastern Silesia that was shown to be conspecific with R. subgothicus, a here lectotypified, formerly overlooked species; (3) a new species from the southern part of the Czech Republic and northern Austria, R. lobifolius, which was included by earlier authors in R. gothicus s. str.; and (4) another new species, R. scarbantinus, which is common in the eastern foothills of the Alps. Flow cytometric measurements confirmed that all the here circumscribed species of the R. gothicus group are tetraploid. We provided a taxonomic and nomenclatural discussion, revised morphological description, ecological characteristics and distribution maps for the species studied.
The morphological variation and cytotype diversity were investigated among Central European populations traditionally recognized as Dactylorhiza fuchsii, recently incorporated in D. maculata s.l. Flow cytometry was employed to assess the ploidy levels of 738 individuals from 77 localities and multivariate morphometrics for a total of 531 individuals from 27 localities. Three ploidy levels were found: diploid (2n = 2x = 40), DNA-triploid and tetraploid (2n = 4x = 80). Whereas diploids and tetraploids often occurred as pure-cytotype populations, individuals of DNA-triploids always co-occurred with at least one of the other cytotypes. Qualitative morphological traits were inferred to be the most important drivers of morphological variation among the investigated plants, with the most striking differences in flower colouration and leaf spotting. The combination of morphological and cytological characters enabled to delimit two separate groups of populations. The first corresponded to D. maculata subsp. fuchsii with morphologically indistinguishable diploid, DNA-triploid and tetraploid individuals, sometimes occurring in mixed-ploidy populations. A complex geographical pattern of cytotype distributions was observed, with diploids scatteredly occurring throughout Central Europe except for Bohemian Massif, which was dominated by tetraploids. The other group of populations represented newly described in this study D. maculata subsp. sooana, subsp. nova, morphologically well-defined and strictly diploid taxon with a restricted geographical range, occurring in the Western Carpathians. A new combination for a hybrid taxon D. × dinglensis nothosubsp. smitakii, comb. nova (= D. maculata subsp. sooana × D. majalis subsp. majalis), was also proposed.
The genus Ficaria is a taxonomically intricate group in which polyploidization and hybridization contribute to taxa diversification. In central Europe, populations of diploid F. calthifolia and tetraploid F. versa subsp. versa occasionally come into contact, which results in an interspecific triploid hybrid, as recently demonstrated using molecular markers, genome size estimation and experimental crossing. In this study, we aimed to estimate the frequency and distribution of the triploid hybrid in central Europe, to identify those phenotypic traits that can be used to discriminate between hybrid and parental taxa and compare the phenology, pollen viability, sexual and asexual reproduction and niche differentiation of the triploid hybrid and parental taxa. Flow-cytometry analyses of 1171 individuals sampled from 67 localities revealed that triploid hybrids were regularly found at 89% of the sites sampled where there were sympatric populations of both parental taxa, with a mean percentage of 19.4% occurring at sites where there was a mixture of cytotypes. No pure hybrid population was found. The hybrids were intermediate between the parental taxa in most morphological characters and did not show any novel morphological characters. The characters that best differentiate the taxonomic groups were the production of axillary bulbils, aborted and well-developed achenes, leaf shape and plant habit. Almost all hybrids were sexually sterile: the pollen viability was considerably reduced (mean 8.4%) and on average 98% of the achenes were aborted. Local dispersal of the hybrids is possible through the production of bulbils in the leaf axils, although the number of bulbils per node and the mean weight of one bulbil were two and three times lower in the hybrid than in F. verna subsp. versa. The hybrid niche was intermediate between those of the parental taxa but shifted slightly towards that of F. yenta subsp. verna. In addition, the taxonomic and nomenclatural treatment of the hybrid, which is described as Ficaria xsellii Duchoslav, Popelka et Travn., as well as a key for identifying the central European taxa of Ficaria. are presented.
The genus Ficaria is a taxonomically intricate group in which polyploidization and hybridization contribute to taxa diversification. In central Europe, populations of diploid F. calthifolia and tetraploid F. verna subsp. verna occasionally come into contact, which results in an interspecific triploid hybrid, as recently demonstrated using molecular markers, genome size estimation and experimental crossing. In this study, we aimed to estimate the frequency and distribution of the triploid hybrid in central Europe, to identify those phenotypic traits that can be used to discriminate between hybrid and parental taxa and compare the phenology, pollen viability, sexual and asexual reproduction and niche differentiation of the triploid hybrid and parental taxa. Flowcytometry analyses of 1171 individuals sampled from 67 localities revealed that triploid hybrids were regularly found at 89% of the sites sampled where there were sympatric populations of both parental taxa, with a mean percentage of 19.4% occurring at sites where there was a mixture of cytotypes. No pure hybrid population was found. The hybrids were intermediate between the parental taxa in most morphological characters and did not show any novel morphological characters. The characters that best differentiate the taxonomic groups were the production of axillary bulbils, aborted and well-developed achenes, leaf shape and plant habit. Almost all hybrids were sexually sterile: the pollen viability was considerably reduced (mean 8.4%) and on average 98% of the achenes were aborted. Local dispersal of the hybrids is possible through the production of bulbils in the leaf axils, although the number of bulbils per node and the mean weight of one bulbil were two and three times lower in the hybrid than in F. verna subsp. verna. The hybrid niche was intermediate between those of the parental taxa but shifted slightly towards that of F. verna subsp. verna. In addition, the taxonomic and nomenclatural treatment of the hybrid, which is described as Ficaria ×sellii Duchoslav, Popelka et Trávn., as well as a key for identifying the central European taxa of Ficaria, are presented.
The paper presents the distribution of Taraxacum bibulum and Taraxacum ranunculus, two species of sect. Palustria new for the Polish flora. The characteristics of the habitat at Polish locations and the taxonomic description and distinguishing features from similar species are given.
In this study we revised apomictic taxa of Rubus ser. Sylvatici occurring in the Pannonian Basin and adjacent regions (i.e. the northern Balkans. Carpathians, and eastern and southern edges of the Alps). The first modern taxonomic studies on this series in the area studied were carried out by W. Maurer (from the 1960s), who described three new species, R. juennensis. R. salzmannii and R. venoms, disentangled the position of R. ferox and presented the first reliable data on R. macrophyllus. A further species, R. wimmerianus. was recorded by various authors in the 1990s in the northern part of the area. Based on comprehensive herbarium revisions, incl. assessment of the original material for several names, and recent field studies, we confirmed that the species in ser. Sylvatici described by Maurer are well-defined and taxonomically well-positioned. In addition, we describe here a widely distributed new species, R. maureri, occurring from Italy to Hungary, that was earlier confused with R. macrophyllus. We present the full synonymy (incl. necessary typifications) and an updated distribution map for R. ferox. and broaden the knowledge on the taxonomic position, chomlogy and ecology of R. juennensis, R. macrophyllus. R. salzmannii, R. venosus and R. wimmerianus. An extended morphological characterization, iconography (pen-drawings, photographs of living plants and type specimens) for most of the species studied are provided. Finally, we discuss the taxonomic position of R. angustipaniculatus and R. solvensis. which were placed by some authors in the ser. Sylvatici, and support keeping the first species in ser. Rhamnifolii and the second in ser. Vestiti.