The article presents an actual checklist of Oxytropis (Fabaceae) of the flora of Uzbekistan compiled on the basis of authors' field observations, critically revised literature, herbarium collections and occurrence records from online platforms. The synopsis includes 62 species, 51 of them are endemic to Mountain Middle Asian floristic province, including 16 endemics of Western Tian Shan and 17 endemics of Pamir-Alay; eight species are national endemics, six are listed in the Red Data Book of Uzbekistan. Three species (O. baldshuanica, O. ovczinnikovii and O. trajectorum) are reported as new for Uzbekistan, O. submutica is a novelty for the flora of Kyrgyzstan. Distribution maps, species richness and collection density map based on all known records are provided; distribution patterns of Oxytropis species in Uzbekistan within phytochoria, mountain ranges and vertical zones are analyzed. Hissar Range in Pamir-Alay and Chatkal Range in Western Tian Shan are identified as "hot spots" of Oxytropis diversity in Uzbekistan.
Nucleotide polymorphism of the psbA–trnH, trnL–trnF, and trnS–trnG intergenic spacers of chloroplast DNA was analyzed to assess the genetic diversity and demographic dynamics of endemic Oxytropis species of the section Orobia in two speciation centers, Southern Siberia and northeast Asia. Of the 34 studied populations of endemic species, five were monomorphic. These populations were found only in the species from northeast Asia. Eighteen populations (52.9
The cosmopolitan genus Hedysarum L. (Fabaceae) is divided into sections Hedysarum, Stracheya, and Multicaulia. This genus includes many valuable medicinal, melliferous, and forage species. The species taxonomy and genome relationships within the sections are still unclear. We examined intra- and interspecific diversity in the section (sect.) Hedysarum based on repeatome analyses using NGS data, bioinformatic technologies, and chromosome FISH mapping of 35S rDNA, 5S rDNA, and the identified satellite DNA families (satDNAs). A comparison of repeatomes of H. alpinum, H. theinum, and H. flavescens revealed differences in their composition. However, similarity in sequences of most satDNAs indicated a close relationship between genomes within sect. Hedysarum. New effective satDNA chromosomal markers were detected, which is important for karyotype analyses within Hedysarum. Intra- and interspecific variability in the chromosomal distribution patterns of the studied markers were revealed, and species karyograms were constructed. These results provided new insight into the karyotype structures and genomic diversity within sect. Hedysarum, clarified the systematic position of H. sachalinense and H. arcticum, and confirmed the distant genomic relationships between species from sections Hedysarum and Multicaulia. Our findings are important for further comparative genome studies within the genus Hedysarum.
A comparative analysis of genomes of six Hedysarum species was carried out using the rapid genomic in situ hybridization (rapidGISH) technique, which makes it possible to identify common highly repetitive DNA sequences, and also shows the patterns of their distribution on chromosomes of the studied species. Dispersed localization of genomic DNA of H. flavescens (sect. Hedysarum) was found on chromosomes of H. alpinum and H. theinum (both belonged to sect. Hedysarum). On chromosomes of H. neglectum from the section Hedysarum, clustered hybridization signals of genomic DNA of H. flavescens were observed. The localization of H. alpinum genomic DNA was found along the all chromosomes of H. flavescens and H. theinum. On chromosomes of H. grandiflorum and H. dahuricum (both from sect. Multicaulia), small clustered and dispersed hybridization signals of genomic DNA of H. flavescens and H. alpinum were found. Thus, in the studied species, common highly repetitive DNA sequences of different types of their chromosome organization were revealed. Our findings indicate the presence of common origin of the species from the sections Hedysarum and Multicaulia with different degree of their relationship.
Genetic diversity and divergence of closely related species Oxytropis strobilacea,O. adamsiana, and O. vassilczenkoi series Strobilacei section Orobia (Fabaceae) from Asian Russia were studied based on nucleotide polymorphism of cpDNA intergenic spacers psbA-trnH,trnL-trnF, and trnS-trnG, as well as ITS nrDNA. Most populations are characterized by a moderate or high level of chloroplast genetic diversity (h varies from 0.600 to 1.000). In total, 65 chlorotypes have been identified, no shared chlorotypes were found in the taxa, which confirms the status of O. vassilczenkoi as a separate species. Two phyletic lineages have been identified in O. strobilacea, which indicates ongoing intensive diversification processes. One of the seven ITS ribotypes is shared for the three species, which is probably due to their common origin and relatively recent divergence.
The section Multicaulia is the largest clade in the genus Hedysarum L. (Fabaceae). Representatives of the sect. Multicaulia are valuable plants used for medicinal and fodder purposes. The taxonomy and phylogeny of the sect. Multicaulia are still ambiguous. To clarify the species relationships within sect. Multicaulia, we, for the first time, explored repeatomes of H. grandiflorum Pall., H. zundukii Peschkova, and H. dahuricum Turcz. using next-generation sequencing technologies and a subsequent bioinformatic analysis by RepeatExplorer/TAREAN pipelines. The comparative repeatome analysis showed that mobile elements made up 20-24% (Class I) and about 2-2.5% (Class II) of their repetitive DNAs. The amount of ribosomal DNA varied from 1 to 2.6%, and the content of satellite DNA ranged from 2.7 to 5.1%. For each species, five high confident putative tandem DNA repeats and 5-10 low confident putative DNA repeats were identified. According to BLAST, these repeats demonstrated high sequence similarity within the studied species. FISH-based mapping of 35S rDNA, 5S rDNA, and satDNAs made it possible to detect new effective molecular chromosome markers for Hedysarum species and construct the species karyograms. Comparison of the patterns of satDNA localization on chromosomes of the studied species allowed us to assess genome diversity within the sect. Multicaulia. In all studied species, we revealed intra- and interspecific variabilities in patterns of the chromosomal distribution of molecular chromosome markers. In H. gmelinii Ledeb. and H. setigerum Turcz. ex Fisch. et Meyer, similar subgenomes were detected, which confirmed the polyploid status of their genomes. Our findings demonstrated a close genomic relationship among six studied species indicating their common origin and confirmed the taxonomic status of H. setigerum as a subspecies of H. gmelinii as well as the validity of combining the sect. Multicaulia and Subacaulia into one sect. Multicaulia.
The dataset providing information on the geographic distribution of Oxytropis species on the territory of Asian Russia is discussed. The data were extracted from different sources including prominent floras and check-lists, Red Data books, published research on congeneric species and authors’ field observations and mainly cover less-studied, remote regions of Russia. The dataset should be of value to applied, basic and theoretical plant biologists and ecologists interested in the Oxytropis species.The dataset includes 5172 distribution records for 143 species and 15 subspecies of genus Oxytropis DC. (Fabaceae Lindl.) in Asian Russia. The dataset fills gaps in the distribution of locoweeds in the study area and contains precise coordinates for many of rare and endemic species.
On the basis of the analysis of the nucleotide polymorphism of the intergenic spacers psbA–trnH, trnL–trnF, and trnS–trnG of chloroplast DNA of the Oxytropis species from Asian Russia, O. tragacanthoides section Hystrix subgenus Tragacanthoxytropis, O. coerulea, O. filiformis, and O. mandshurica sect. Janthina, and O. deflexa and O. glabra sect. Mesogaea subg. Phacoxytropis, it was found that all populations are characterized by high haplotype diversity (h varies from 0.676 to 1.000), except for species of the sect. Mesogaea (h varies from 0 to 0.333). Species-specific markers were found for O. tragacanthoides, O. deflexa, O. glabra, and O. mandshurica, as well as specific markers for the sect. Mesogaea. Reconstruction of the phylogenetic relationships of the chlorotypes of the species of the subg. Phacoxytropis, Tragacanthoxytropis, and Oxytropis showed that the species of the sect. Janthina are combined into one well-supported clade with the species of the subg. Tragacanthoxytropis and Oxytropis, but their relationships remained unresolved. An analysis of the genealogical relationships of the ribotypes of the ITS of nuclear DNA revealed a common ribotype for the species O. tragacanthoides, O. coerulea, O. lanata, O. chankaensis, O. oxyphylla, and O. triphylla, belonging to three subgenera. The revealed genetic affinity with clear morphological differences is characteristic of taxa with a common origin that have experienced relatively recent rapid adaptive radiation. The obtained data on the variability of markers of the nuclear and chloroplast genomes confirm the status of O. coerulea, O. filiformis, and O. mandshurica as three separate species.
The seed production of the rare species Gueldenstaedtia monophylla has been investigated in 19 coenotic populations in the territories of Russia (Central Altai) and Mongolia (the north-western part). G. monophylla reproduces exclusively by seeds and is characterized by the low number and density of the plants in the populations. Its populations have a high proportion of generative plants: from 31 to 84% from the total number of the plants. The plants of G. monophylla have a small number of generative shoots, varying from 1 to 6 per plant. The number of flowers on the plants is low, too, and varies in the populations on average from 1.2 to 15.3 per plant. The potential seed production (PSP) varies considerably from 13.5 to 308.6 ovules per plant. The real seed production is much lower, to constitute from 6.2 to 74.5 seeds per plant. The low seed production, fructification that occurs not every year, and scanty populations make the species vulnerable.
The systematic knowledge on the genus Hedysarum L. (Fabaceae: Hedysareae) is still incomplete. The species from the section Hedysarum are valuable forage and medicinal resources. For eight Hedysarum species, we constructed the integrated schematic map of their distribution within Eurasia based on currently available scattered data. For the first time, we performed cytogenomic characterization of twenty accessions covering eight species for evaluating genomic diversity and relationships within the section Hedysarum. Based on the intra- and interspecific variability of chromosomes bearing 45S and 5S rDNA clusters, four main karyotype groups were detected in the studied accessions: (1) H. arcticum, H. austrosibiricum, H. flavescens, H. hedysaroides, and H. theinum (one chromosome pair with 45S rDNA and one pair bearing 5S rDNA); (2) H. alpinum and one accession of H. hedysaroides (one chromosome pair with 45S rDNA and two pairs bearing 5S rDNA); (3) H. caucasicum (one chromosome pair with 45S rDNA and one chromosome pair bearing 5S rDNA and 45S rDNA); (4) H. neglectum (two pairs with 45S rDNA and one pair bearing 5S rDNA). The species-specific chromosomal markers detected in karyotypes of H. alpinum, H. caucasicum, and H. neglectum can be useful in taxonomic studies of this section.
For the first time, a comparative karyotype analysis of closely related species Hedysarum gmelinii andH. setigerum (Hedysarum section Multicaulia) grown in Southern Siberia, has been performed by molecular cytogeneticmarkers. Chromosome numbers in karyotypes of these species were specified – 2n = 4х = 32. In some accessions, additionalB chromosomes were revealed. FISH analyses indicated high similarities in chromosome morphology and also patternsof chromosomal distributions of 45S and 5S rDNA clusters in karyotypes of H. gmelinii and H. setigerum, which confirmsthe close relationship between their genomes.
For the first time, chromosomal polymorphism in karyotypes of three species from the section Hedysarum (= syn. Gamotion) of the genus Hedysarum L. (Fabaceae) grown in Southern Siberia has been studied with the useof molecular cytogenetic markers. This comparative molecular cytogenetic analysis revealed high similarity in morphology of chromosomes in H. alpinum L., H. austrosibiricum B. Fedtsch. and H. theinum Krasnob. as well as in patterns ofdistribution of 45S and 5S rDNA loci in their karyotypes confirming their close relationships. Considerable intra-specificpolymorphism on 45S rDNA chromosome localization was detected in H. theinum. In karyotype of H. alpinum, unlikethe other two species, two chromosome pairs bearing 5S rDNA locus were observed which could be used as additionalspecies-specific markers.
Modern botanical studies revealing patterns of plant species distribution are based on analysis of big datasets. Despite publishing many maps of diversity and species richness on the global scale and for huge biogeographic regions of the world, the territories of Northern and Central Asia remain poorly studied. We elaborated a special database, including distribution of 19 Oxytropis species of the section Xerobia with 1353 localitites (See Fig. 1). For all species, we analyzed their whole distribution range, including data from Kazakhstan, Mongolia and China. Species distribution was detected according to the main foreign and Russian herbaria, online databases, field data and relevés. Species distribution modeling was performed using Maxent 3.3.3k with MIROC-ESM model in resolution of 2.5 arc-minutes. 19 BIOCLIM and 18 ENVIREM variables were analyzed. Past climate change was evaluated using ENVIREM variables for the Mid-Holocene (ca. 6.000 yr. BP) and the Last Glacial Maximum (ca. 22.000 yr. BP). Future distribution modeling was carried out basing on different climatic scenarios, according to IPCC AR5: RCP8.5, RCP2.6 and RCP6.0. Species distribution from the section Xerobia mostly occupied the territory of Central Asia (See Fig. 1). Few species, such as Oxytropis grandiflora (Pall.) DC. and O. leptophylla (Pall.) DC., were mostly found in the western part of Xerobia section distribution on the territory of Zabaykal’skiy region of Russia, Eastern province of Mongolia and Inner Mongolia province of China. The most part of Xerobia species have isolated distribution and often occupy specific habitats. In such case, using SDM with only bioclimatic variables for local endemic species is pointless. So, we chose species Oxytropis ampullata (Pall.) Pers. (See Fig. 2) with Central Asian distribution and O. grandiflora with Manchuro-Dahurian distribution for modeling (See Fig. 3A). The selected species differ in their ecology: O. ampullata is a mountainous species, whereas most habitats for O. grandiflora are river valleys and mid-mountainous regions. Our analysis showed that ENVIREM variables provide more correct modeling results than BIOCLIM variables (See Fig. 2). Predictive maps on the basis of BIOCLIM variables showed wide potential distribution for O. ampullata, which does not correspond well to the species ecology. The main habitats for this species are such mountainous regions as the Khangai mountains, the Russian and the Mongolian Altai mountains, the Dzhungarian mountains, and the Tarbagatai ridge. Additionally, modeling showed potential distribution for the species in the Selenga river valley. Modern distribution of O. grandiflora was studied quite well; suitable habitats with new localities for the species can be found in the Khentii mountains (See Fig. 3A). The determinants for O. ampullata are mean annual temperature, isothermality and potential evapotranspiration (PET) of the driest and coldest quarter (See Table 1). PET parameters in the driest and the coldest time of the year have the key meaning because in arid conditions plants receive the main portion of moisture in the colder period when the evaporation is not intense, also it is important to conserve the moisture during the dry season. Distribution of O. grandiflora is limited by temperature and precipitation seasonality, temperature annual range, PET seasonality, and PET of the driest quarter (See Table 1). Determinants for the species with Central-Asian distribution O. ampullata are connected with temperature variables, whereas for Manchuro-Daurian species O. grandiflora precipitation matters (See Table 1 and Fig. 3A). The key factors for modern distribution of the studied Xerobia species are mean monthly potential evapotranspiration of the driest quarter, continentality index and climatic moisture index (See Fig. 3B). All these variables were determinants for the mid-Holocene and the Last Glacial Maximum (See Table 2 and Fig. 4), which might give evidence of relatively stable environmental conditions in the studied region. Central Asia has not been severely affected by glaciation as more northern latitudes and climate conditions on that territory were relatively stable during a long period. Modeling for the past climate showed a wider distribution for Xerobia species in the north-west during the Last Glacial Maximum and future shrinking during the Mid-Holocene till modern time (See Fig. 4). The north-eastern territories, such as Zabaykal’skiy region of Russia and, partially, the central part of Siberia, are characterized by a wider distribution under modern climate conditions. Species habitats of that territory are mostly confined with mountains. It is consistent with previous studies that described Southern Siberia as one of the centers of speciation for the genus Oxytropis. This region has now high Oxytropis species richness with a great number of endemics.Predictive maps for different climate scenarios reveal insignificant changes in distribution of the section Xerobia, even for the maximum climate warming (RCP8.5 scenario) (See Fig.5). Under predicted climate change, potential habitats in the southwest and in the north-east of Xerobia distribution, as well as a slight shrinking in the south-east can be observed in the future.
The seed productivity of endemic species of the genus Hedysarum L. in southern Siberia was studied. It is established that the reproduction of these species is carried out only by seed. Flowering occurs in June, the number of flowers in the inflorescence ranges from 12.3 to 19.1. The efficiency of fruit formation (% of the set beans from the number of flowers) differs significantly: the maximum values are marked for H. austrosibiricum, H. turczaninovii and H. sangilense (90.7-74.1 %), the minimum values are marked for the rare H. minussinense, H. chaiyrakanicum, H. zundukii (35.3 – 47.2 %). A decrease in the number of seeds that are set compared to the number of ovules, caused by various reasons, leads to a significant decrease in the RSP compared to the PSP. The percentage of semenification in the studied species is highest in the endemics H. austrosibiricum, H. sangilense and H. turczaninovii (56.545.1 %), the minimum – in the rare H. zundukii, H. minussinense and H. chaiyrakanicum (9.8-36.5 %).
The article presents the results of the analysis of the demographic structure of ten coenotic populations of Oxytropis chakassiensis in near-Yenissei steppes (Khakassia and Krasnoyarsk krai). Most part of studied species coenopopulations are not complete and definitive, normal, mature, have a bimodal developmental spectrum with peaks at g1 (v)- and g3-individuals. Changes in the structure of the ontogenetic spectrum (increasing the share of young plants) depend mainly on the environmental conditions and the degree of human disturbance of habitats: rocky slopes and decrease in pasture digression create more favorable conditions for the survival of juveniles. The ontogeny of O. chakassiensis (the rare species of near-Yenissei steppes) was studied and its life- form was described. Four periods and nine ontogenic stages were distinguished. The results of our study showed that O. chakassiensisis was a petrophyte steppe species with strict ecological and cenotic propensity to petrophytic steppe with the prevalence Koeleria cristata, Festuca valesiaca, Arctogeron gramineum, Hedysarum turczaninovii, Alyssum obovatum and Thymus minussinensis. Due to long generative period O. chakassiensis is successfully renewed and is stable during long periods of extreme conditions of petrophytic steppes with high pasture digression.
The anatomical features of the structure of the leaf blade were used to trace the adaptation of species of the Hedysarum L. section Gamotion Basin. to different growing conditions in South Siberia. On the example of consanguineum DC. and 11. rhenium Krasnob, it was revealed that in high gradient habitats heliomorphic features were enlarged: size of plants and leaves was reduced: the thickness of the lamina, ply palisade mesophylk height of palisade and epidermal cells, number of stomata and total idioblast were increased. It was revealed that the anatomical structure of the leaf species of the Gamotion section occupying different ecological niches in the mountains and intermountain basins of southern Siberia had undergone similar changes in the process of adaptation to modern conditions of growth. The plants of H. neglectum Ledeb. from the forest belt and of IL afpinum L. from the foothill plains in drier habitats had the same changes as the Alpine species in the high-altitude gradient of habitats. The increase in the thickness and volume of the leaf blade, the total volume of idioblasts, the number of rows and height of palisade cells in the structure of the leaf in species of the Iledysarum section Gamotion can be considered as important biological adaptations to extreme conditions of growth in South Siberia.
В работе представлены результаты изучения кариотипов редких и эндемичных видов Astragalus chorinensis Bunge, Oxytropis glandulosa Turcz., O. stukovii Palibin, O. varlakovii Serg. На основе полученных данных представлены идиограммы хромосом и формулы кариотипов. Исследование показало, что у A. chorinensis соматическое число хромосом 2n = 16. Хромосомная формула 2n = 2m + 14sm. Средняя суммарная длина гаплоидного набора составляет 40,24 мкм. Размеры хромосом варьируют от 4,29 до 6,31 ± 0,06 мкм. A. chorinensis характеризуется асимметричным кариотипом с преобладанием субметацентрических хромосом. Диплоидный набор у O. glandulosa 2n = 16, хромосомная формула 2n = 16 = 4m + 12sm. Хромосомы достаточно однородны по размеру, средняя длина составила от 1,56 ± 0,05 до 1,93 ± 0,09 мкм. Суммарная длина гаплоидного набора хромосом составила 33,63 мкм. Для O. stukovii установлена хромосомная формула 2n = 16 = 6m + 10sm. Общая длина гаплоидного набора (CL) составила 33,74 мкм. Длина хромосом колеблется от 3,75 ± 0,10 мкм до 4,93 ± 0,14 мкм. Хромосомы O. varlakovii относятся к метацентрическому и субметацентрическому типу. Хромосомная формула 2n = 32 = 4m + 28sm. Размеры хромосом варьируют от 0,74 ± 0,01 до 1,56 ± 0,15 мкм. Суммарная длина гаплоидного набора составила 33,74 мкм. O. stukovii, O. glandulosa и O. varlakovii характеризуются симметричным кариотипом с преобладанием хромосом метацентрического типа.
The influence of climatic factors, e.g., low temperature, on the phytochemical composition and bioactivity of the arctic plant Dracocephalum palmatum Steph. ax Willd. (palmate dragonhead), a traditional food and medical herb of Northern Siberia, was investigated. D. palmatum seedlings were grown in a greenhouse experiment at normal (20 °C, NT) and low (1 °C, LT) temperature levels and five groups of components that were lipophilic and hydrophilic in nature were characterized. The analyses indicated that D. palmatum under NT demonstrates high content of photosynthetic pigments, specific fatty acid (FA) profile with domination of saturated FA (53.3%) and the essential oil with trans-pinocamphone as a main component (37.9%). Phenolic compounds were identified using a combination of high performance liquid chromatography with diode array detection and electrospray ionization mass-spectrometric detection (HPLC-DAD-ESI-MS) techniques, as well as free carbohydrates and water soluble polysaccharides. For the first time, it was established that the cold acclimation of D. palmatum seedlings resulted in various changes in physiological and biochemical parameters such as membrane permeability, photosynthetic potential, membrane fluidity, leaf surface secretory function, reactive oxygen species–antioxidant balance, osmoregulator content and cell wall polymers. In brief, results showed that the adaptive strategy of D. palmatum under LT was realized on the accumulation of membrane or surface components with more fluid properties (unsaturated FA and essential oils), antioxidants (phenolic compounds and enzymes), osmoprotectants (free sugars) and cell wall components (polysaccharides). In addition, the occurrence of unusual flavonoids including two new isomeric malonyl esters of eriodictyol-7-O-glucoside was found in LT samples. Data thus obtained allow improving our understanding of ecophysiological mechanisms of cold adaptation of arctic plants.
В работе представлены результаты изучения кариотипов редких и эндемичных видов Astragalus chorinensis Bunge, Oxytropis glandulosa Turcz., O. stukovii Palibin, O. varlakovii Serg. На основе полученных данных представлены идиограммы хромосом и формулы кариотипов. Исследование показало, что у A. chorinensis соматическое число хромосом 2n = 16. Хромосомная формула 2n = 2m + 14sm. Средняя суммарная длина гаплоидного набора составляет 40,24 мкм. Размеры хромосом варьируют от 4,29 до 6,31 ± 0,06 мкм. A. chorinensis характеризуется асимметричным кариотипом с преобладанием субметацентрических хромосом. Диплоидный набор у O. glandulosa 2n = 16, хромосомная формула 2n = 16 = 4m + 12sm. Хромосомы достаточно однородны по размеру, средняя длина составила от 1,56 ± 0,05 до 1,93 ± 0,09 мкм. Суммарная длина гаплоидного набора хромосом составила 33,63 мкм. Для O. stukovii установлена хромосомная формула 2n = 16 = 6m + 10sm. Общая длина гаплоидного набора (CL) составила 33,74 мкм. Длина хромосом колеблется от 3,75 ± 0,10 мкм до 4,93 ± 0,14 мкм. Хромосомы O. varlakovii относятся к метацентрическому и субметацентрическому типу. Хромосомная формула 2n = 32 = 4m + 28sm. Размеры хромосом варьируют от 0,74 ± 0,01 до 1,56 ± 0,15 мкм. Суммарная длина гаплоидного набора составила 33,74 мкм. O. stukovii, O. glandulosa и O. varlakovii характеризуются симметричным кариотипом с преобладанием хромосом метацентрического типа.