Hybrids as mirrors of the past: genomic footprints reveal spatio-temporal dynamics and extinction risk of alpine extremophytes in the mountains of Central Asia.

Hybridization is one of the key processes shaping lineage diversification, particularly in regions that experienced strong climate oscillations. The alpine biome with its rich history of glacial-interglacial cycles and complex patterns of species distribution shifts offers an excellent system to investigate the impact of gene flow on population dynamics and speciation, important issues for evolutionary biology and biodiversity conservation. In this study, we combined genomic data (DArTseq), chloroplast markers, and morphology to examine phylogenetic relationships and the permeability of species boundaries and their evolutionary outcomes among the alpine extremophilic species of Puccinellia (Poaceae) in the Pamir Mountains, a part of the Mountains of Central Asia biodiversity hotspot. We determined the occurrence of interspecific hybrids between P. himalaica and P. pamirica, which demonstrated almost symmetric ancestry from their parental species and did not show signals of introgression. According to our integrative revision, the natural hybrids between P. himalaica and P. pamirica should be classified as Puccinellia ×vachanica (pro species). Using approximate Bayesian computation for population history inference, we uncovered that P. himalaica hybridized with P. pamirica independently in multiple localities over the Holocene. Hybrids inherited the fine-scale genetic structure from their parental species, which developed these patterns earlier, during the Late Pleistocene. Hybridization had different consequences for the involved parental lineages, likely playing an important role in a continuing decline of P. himalaica in the Pamir Mountains over the Holocene. Our results show that P. himalaica should be considered a critically endangered species in the Pamir Mountains and could also be retreating across its entire range of distribution in High Mountain Asia. Using a comparative phylogeographic framework, we revealed the risk of extinction of a cold-adapted alpine species in a global biodiversity hotspot. This study highlights that genomics could unravel diversity trends under climate change and provides valuable evidence for conservation management.

Insight into the phylogeny and responses of species from the genus Sergia (Campanulaceae) to the climate changes predicted for the Mountains of Central Asia (a world biodiversity hotspot).

BACKGROUND: Together with other elevated areas, the Mountains of Central Asia are significantly threatened by ongoing climate change. The presence of refuges during the glaciations makes the region extremely rich in species, especially endemic ones. However, the limited potential for colonisation of other habitats makes rocky-related species with 'island-like' distribution, particularly vulnerable to climate change. To understand the processes underlying species response to climate warming, we assessed differences in ecological niches and phylogenetic relationship of two geographically disjunctive alpine species belonging to the genus Sergia. The taxa are considered Tertiary relicts, endemic to the Tian Shan and Pamir-Alai Mountains. To illustrate range dynamics and differences in occupied niches of Sergia species, we used Ecological Niche Modelling of current and future distribution. Whereas, to reconstruct the phylogenetic relationship within and between Sergia and other related Campanulaceae species from the region we used molecular data (ITS, cpDNA, DArTseq-derived SNPs). RESULTS: The results reveal that the genus Sergia is a polyphyletic group, and its representatives differ geographically, ecologically and genetically. Both S. regelii and S. sewerzowii constitute a common clade with Asyneuma group, however, S. sewerzowii is more closely related to Campanula alberti (a species that has never previously been considered closely related to the genus Asyneuma or Sergia) than to S. regelii. Sergia sewerzowii is adapted to lower elevations with higher temperatures, while S. regelii prefers higher elevations with lower temperatures. The future distribution models demonstrate a dramatic loss of S. regelii range with a shift to suitable habitats in higher elevations, while the potential range of S. sewerzowii increases and shifts to the north. CONCLUSIONS: This study shows that S. regelii and S. sewerzowii have a long and independent evolution history. Sergia regelii and S. sewerzowii significantly differ in realised niches. These differences are mirrored in the response of the studied endemics to future climate warming. As suitable habitats shrink, rapid changes in distribution can lead to species' range loss, which is also directly related to declines in genetic variability. The outcomes of this paper will help to more precisely assess the impact of climate changes on rocky-related plant species found in this world's biodiversity hotspot.

Hybridization and introgression events in cooccurring populations of closely related grasses (Poaceae: Stipa) in high mountain steppes of Central Asia.

Stipa is a genus comprising ca. 150 species found in warm temperate regions of the Old World and around 30% of its representatives are of hybrid origin. In this study, using integrative taxonomy approach, we tested the hypothesis that hybridization and introgression are the explanations of the morphological intermediacy in species belonging to Stipa sect. Smirnovia, one of the species-rich sections in the mountains of Central Asia. Two novel nothospecies, S. magnifica × S. caucasica subsp. nikolai and S. lingua × S. caucasica subsp. nikolai, were identified based on a combination of morphological characters and SNPs markers. SNPs marker revealed that all S. lingua × S. caucasica samples were F1 hybrids, whereas most of S. magnifica × S. caucasica samples were backcross hybrids. Furthermore, the above mentioned hybrids exhibit transgressive morphological characters to each of their parental species. These findings have implications for understanding the process of hybridization in the genus Stipa, particularly in the sect. Smirnovia. As a taxonomic conclusion, we describe the two new nothospecies S. × muksuensis (from Tajikistan) and S. × ochyrae (from Kyrgyzstan) and present an identification key to species morphologically similar to the taxa mentioned above.

Alpine Extremophytes in Evolutionary Turmoil: Complex Diversification Patterns and Demographic Responses of a Halophilic Grass in a Central Asian Biodiversity Hotspot.

Diversification and demographic responses are key processes shaping species evolutionary history. Yet we still lack a full understanding of ecological mechanisms that shape genetic diversity at different spatial scales upon rapid environmental changes. In this study, we examined genetic differentiation in an extremophilic grass Puccinellia pamirica and factors affecting its population dynamics among the occupied hypersaline alpine wetlands on the arid Pamir Plateau in Central Asia. Using genomic data, we found evidence of fine-scale population structure and gene flow among the localities established across the high-elevation plateau as well as fingerprints of historical demographic expansion. We showed that an increase in the effective population size could coincide with the Last Glacial Period, which was followed by the species demographic decline during the Holocene. Geographic distance plays a vital role in shaping spatial genetic structure of P. pamirica alongside with isolation-by-environment and habitat fragmentation. Our results highlight a complex history of divergence and gene flow in this species-poor alpine region during the Late Quaternary. We demonstrate that regional climate specificity and a shortage of non-climate data largely impede predictions of future range changes of the alpine extremophile using ecological niche modelling. This study emphasises the importance of fine-scale environmental heterogeneity for population dynamics and species distribution shifts.

Taxonomy and distribution of Taraxacum sect. Erythrosperma (Asteraceae) in Poland.

The dandelions from Taraxacumsect.Erythrosperma are taxonomically well distinguished and ecologically restricted to warm and sunlit habitats of steppes, dry and sandy grasslands, and distributed in temperate regions of Europe and Central Asia, with some being introduced to North America. Despite the long tradition of botanical research, the taxonomy and distribution of dandelions of T.sect.Erythrosperma is still underexplored in central Europe. In this paper, by combining traditional taxonomic studies supported by micromorphological, molecular and flow cytometry analyses as well as potential distribution modelling we shed light on taxonomical and phylogenetical relationships between members of T.sect.Erythrosperma in Poland. We also provide an identification key, species-checklist, detailed descriptions of morphology and occupated habitats as well as distribution maps for 14 Polish erythrosperms (T.bellicum, T.brachyglossum, T.cristatum, T.danubium, T.disseminatum, T.dissimile, T.lacistophyllum, T.parnassicum, T.plumbeum, T.proximum, T.sandomiriense, T.scanicum, T.tenuilobum, T.tortilobum). Finally, conservation assessments performed using the IUCN method and threat categories for all the examined species are proposed.

Temperature and Precipitation More Than Tree Cover Affect the Distribution Patterns of Epiphytic Mosses within the Orthotrichaceae Family in China and Adjacent Areas.

Epiphytes, including vascular and non-vascular, constitute a large part of global plant biodiversity. Distribution of obligatory epiphytic bryophytes results from climate and local habitat conditions. The most important epiphytic bryophytes and at the same time poorly investigated and taxonomically problematic ones belong to the family Orthotrichaceae. Epiphytic mosses are also ideal organisms for species modelling, because of having no roots, they are highly dependent on external environmental conditions. For this purpose, we used the ecological niche modelling approach to define their potential distribution in China and adjacent areas and explore factors that shape this distribution. We used 617 occurrence records of 23 species from six genera within the Orthotrichaceae family. Our results suggest that the distribution of members of the Orthotrichaceae family is predominantly affected by bioclimatic variables, especially bio10 (mean temperature of the warmest quarter), bio15 (precipitation seasonality), bio18 (precipitation of the warmest quarter), bio19 (precipitation of the coldest quarter), bio9 (mean temperature of the driest quarter), and bio2 (mean diurnal range). However, the distribution of particular genera is ruled by a different set of those variables. The distribution of two genera (Leratia and Ulota) is also highly influenced by land cover (especially mixed/other trees), whereas human footprint shows a moderate contribution to models of three genera (Lewinskya, Orthotrichum, Nyholmiella). Based on the occupied climatic niche and distribution patterns, representatives of the studied family are divided into two groups. The 'western-montane group' is characterised by lower temperatures and lower precipitation whereas the 'eastern-lowland' group' by more humid and warmer conditions.

Effect of Rudbeckia laciniata invasion on soil seed banks of different types of meadow communities.

In the last decades, biological invasions become the main driver of biodiversity loss. The changes can be noticed not only in the above-ground diversity but also in the underground, including seed banks of native vegetation. In this study, we focus on Rudbeckia laciniata, a species introduced to many European and Asian countries, to characterize its soil seed bank as well as to answer the question, how the species influenced soil seed banks of meadow plant communities in two types of habitats (fresh and wet), where traditional mowing was abandoned. Within the habitats, we conducted our study on a three-step scale of invasion, from full invasion, through the transition zone to the control zone, where no invasion of the species has been observed so far. The majority of the R. laciniata seeds were located in the surface layer of soil. We detected, that 47% (in fresh meadow) and 56% (wet meadow) of recorded species occurred only in a soil seed bank, and were absent in aboveground vegetation. Emergence of native plants from the soil seed bank is low due to rapid shading of the soil surface by R. laciniata seedlings. However, a short-term seed bank of the species gives hope that returning to regular mowing brings the desired results in its the elimination from vegetation, in a fairly short time.

Typification of five plant names described based on specimens collected by Józef Warszewicz in Central and South America.

Józef Warszewicz (1812-1864) was one of the first Polish naturalists to explore the flora of the tropical New World. During two expeditions to Central and South America (1844-1850 and 1850-1853) he collected and delivered to Europe up to twenty thousand plant specimens. To honour his service and his achievements in plant collections, different taxonomists described more than 100 taxa using the surname Warszewicz, for example in the genus name (Warszewiczia) and the species epithets (warszewiczii, warscewiczii, warszewicziana). Unfortunately, a large part of Warszewicz's collection of plant species deposited in the Berlin Herbarium (B), including many type specimens was destroyed during World War II. During digitisation of herbarium collections preserved in the Herbarium of the Jagiellonian University (KRA), we reviewed more than 650 herbarium sheets with plant specimens collected by Warszewicz and originating from his trips to Central and South America. In this paper, we present the typification of five names of species, described base on Warszewicz's plant material. We select lectotypes for Berberiswarszewiczii, Esenbeckiawarscewiczii, Psammisiaramiflora, Remijiainvolucrata and Rondeletiaorthoneura, and provide data on the presence of 17 specimens (isotypes) representing Esenbeckiacornuta, an extremely rare species, that to date is known only from the type locality in Peru. A list of all Warszewicz's specimens preserved at KRA herbarium is also presented. Additionally, in the result of the revision of syntypes of Berberismultiflora and Rondeletiareflexa we designated here the lectotypes for these taxa.

Resurrection of Stipatremula and taxonomy of the high-alpine species from the Stipapurpurea complex (Poaceae, Pooideae).

Stipapurpurea is a high-alpine species that occurs in cryophilous steppes, semi-deserts and stony slopes, from the Tian Shan and Pamirian Plateau through Qinghai-Xizang Plateau to the Himalayas and is characterised by a great morphological variability. During the revision of specimens of the taxon, we observed that the pattern of this variability is linked to the geographical distribution of the specimens. Numerical analyses (PCA and UPGMA) revealed three groups of OTUs corresponding to three morphotypes within the S.purpurea complex. A set of macro- and micromorphological characters, supported by a map of general distributional ranges, are presented to distinguish each of the three taxa within the complex and we reassess the status of Lasiagrostistremula described by Ruprecht in 1869. As a result, Stipatremula, S.purpurea and S.arenosa were distinguished within the complex. The intermediate characters of S.arenosa may suggest its putative hybrid origin (S.tremula × S.purpurea), whereas the presence of extremely long florets may be an expression of the gigas effect. We propose two new combinations (S.tremula and S.arenosa), describe a new nothospecies (S.×ladakhensis) that originated from hybridisation between S.klimesii and S.purpurea s.l. and designate the lectotype for Ptilagrostissemenovii. An identification key and detailed morphological description of species from the S.purpurea complex are also presented.

Red List of vascular plants of Tajikistan - the core area of the Mountains of Central Asia global biodiversity hotspot.

Central Pamir-Alai, which is located almost entirely within the area of Tajikistan, is one of the world hotspots of biodiversity, harbouring ca. 4,300 species and 1,400 endemic plants. The first application of the IUCN Red List criteria reveals that among all native species occurring in Tajikistan 1,627 taxa (38.11%) are threatened, including 23 extinct (0.54%), 271 (6.34%) critically endangered (CR), 717 (16.79%) endangered (EN) and 639 (14.96%) vulnerable (VU). Globally, 20 taxa are extinct, 711 (16.65%) threatened, including 144 (3.37%) critically endangered, 322 (7.54%) endangered and 245 (5.73%) vulnerable. As we found positive correlation between human density and the number of threatened species, we suspect this indirect factor responsible for the species diversity decline. Extinct or threatened taxa have short blooming periods in spring or early summer, have limited geographical range and inhabit mainly valley bottoms at lower altitudes. Threatened taxa occupy extremely dry or wet habitats, such as deserts, semi-deserts, water reservoirs and fens. The group of threatened plants consists mostly of Central Asian, Indo-Indochinese and Arctic species. Ornamental plants have a higher extinction risk than other plants, but species collected for medicinal reasons and used for forage or food reveal lower retreatment rate. Our assessment fills a gap for important plant area and provides the data for raising the effectiveness of plant diversity conservation.

Development and characterization of microsatellite markers for endangered species Stipa pennata (Poaceae) and their usefulness in intraspecific delimitation.

Stipa pennata (Poaceae), has become a rare and endangered species in Central Europe due habitat loss and fragmentation. This species is characterized by high morphological variability, which has resulted in the description of numerous intraspecific taxa. The aim of present work is to develop microsatellite markers useful in population genetics studies as well as in intraspecific taxonomy of S. pennata s.l. We developed ten microsatellite markers using Illumina high-throughput. Polymorphism at each marker was evaluated using 4-15 individuals from four morphotypes of S. pennata s.l. Seven markers showed polymorphism while three were monomorphic. The number of alleles per locus ranged from 7 to 12, and the observed and expected heterozygosity varied from 0.000 to 1.000 and 0.000 to 0.8670, respectively. Our results confirm that three of four studied morphotypes are genetically distinct. The microsatellite markers developed here will be useful for evaluating levels of genetic diversity and differentiation, to study gene flow, population dynamics and in future conservation studies as well as for intraspecific delimitation of morphologically similar taxa within S. pennata s.l.

Plastid super-barcodes as a tool for species discrimination in feather grasses (Poaceae: Stipa).

Present study was designed to verify which or if any of plastome loci is a hotspot region for mutations and hence might be useful for molecular species identification in feather grasses. 21 newly sequenced complete plastid genomes representing 19 taxa from the genus of Stipa were analyzed in search of the most variable and the most discriminative loci within Stipa. The results showed that the problem with selecting a good barcode locus for feather grasses lies in the very low level of genetic diversity within its plastome. None of the single chloroplast loci is polymorphic enough to play a role of a barcode or a phylogenetic marker for Stipa. The biggest number of taxa was successfully identified by the analysis of 600 bp long DNA fragment comprising a part of rbcL gene, the complete rbcL-rpl23 spacer and a part of rpl23 gene. The effectiveness of multi-locus barcode composed of six best-performing loci for Stipa (ndhH, rpl23, ndhF-rpl32, rpl32-ccsA, psbK-psbI and petA-psbJ) didn't reach 70% of analyzed taxa. The analysis of complete plastome sequences as a super-barcode for Stipa although much more effective, still didn't allow for discrimination of all the analyzed taxa of feather grasses.

Stipa pennata subsp. ceynowae (Poaceae, Pooideae), a new taxon from Central Europe.

Based on numerical analyses of macromorphological characters, scanning electron microscopy observation of leaves and lemma micromorphology, as well as field observations, Stipa pennata subsp. ceynowae was described here as a new taxon from Poland. It differs from the most similar S. pennata subsp. pennata and S. borysthenica mainly by its longer ligules of vegetative shoots. The affinities of this taxon are discussed and a morphological comparison with related species is provided. Illustrations and images of the micromorphological structures, as well as information about its distribution, habitat and conservation status are given.