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Taxonomy and phylogeny of Stipa

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Marcin Nobis
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Understanding species distribution, genetic diversification and evolutionary history is extremely important for mountainous regions with a high diversity of endemic species, which are particularly sensitive to climate change. In this study, we use environmental and molecular data obtained from genome-wide analyses to infer the genetic variability, demographic processes, and response of the cold-adapted, endemic geographical-vicariants Stipa gracilis (distributed in the Tian Shan Mts) and S. zeravshanica (distributed in the western Pamir-Alai Mts) to Quaternary climatic oscillations in a Central Asian mountain biodiversity hotspot. Genomic-based reconstructions of demographic history indicate that the examined endemics presented larger effective population sizes during the Last Glacial Maximum (LGM) period and experienced parallel demographic declines afterward. The results of fastSTRUCTURE analysis revealed three genetic clusters within S. gracilis populations and two within S. zeravshanica. The past distribution models reveals the glacial connectivity of both species, resulting in the detection of an admixture of S. zeravshanica genes in the specimens from the westernmost 'Alaian' population of S. gracilis. Although the occurrence of both species is closely associated with calcareous rocks, the differences in the ranges of the species distributions depend mostly on climatic factors, especially temperature and precipitation. The wider realized ecological niche of S. gracilis allows it to better adapt to global warming and potentially extend its range in the future, while S. zeravshanica, with its narrower niche, is more susceptible to environmental changes and potentially at risk of extinction. The findings will contribute to a better understanding of the factors shaping the distribution and genetic differentiation of mountain endemic species and provide a theoretical basis for their conservation by identifying areas sensitive to climate change in biodiversity hotspots.
Marcin Nobis
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Citation: Nobis M, Krzempek M, Nowak A, Gudkova PD, Klichowska E (2022) Resurrection of Stipa tremula and taxonomy of the high-alpine species from the Stipa purpurea complex (Poaceae, Pooideae). PhytoKeys 196: 21-47. Abstract Stipa purpurea 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 Lasiagrostis tremula described by Ruprecht in 1869. As a result, Stipa tremula, 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 Ptilagrostis semenovii. An identification key and detailed morphological description of species from the S. purpurea complex are also presented.
Marcin Nobis
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Over the past two centuries, Altai krai, located in the southeastern part of Western Siberia, has undergone significant transformation as a result of the virgin lands development. The vast even territories of the krai, formerly occupied by steppes dominated by the feather grass, are now almost completely plowed up. Many types of steppe communities, as well as plant species associated with them, have become rare. This paper presents the results of the taxonomic revision of the genus Stipa. The genus has a high conservation value and seven of twelve feather grass species growing in the studied area are listed in the “Red Book of Altai Krai”. Based on the revision of the herbarium collections stored in ALTB, LE, NS, NSK, TK and the authors’ collections, a checklist of the genus Stipa in Altai krai and an illustrated identification key were compiled. The synopsis for each species contains a nomenclature name, a brief description, a note on taxonomy, flowering period and typical habitats. In addition, it presents the distribution of species in the territory of the krai in the form of an administrative districts list and point maps. A new record of S. sareptana species, new for Altai krai and Western Siberia, was found while the presence of S. baicalensis was not confirmed. Additionally, lectotype of S. praecapillata is also here designated.
Marcin Nobis
added a research item
Hybridisation in the wild between closely related species is a common mechanism of speciation in the plant kingdom and, in particular, in the grass family. Here we show the usefulness of integrative taxonomy as a tool for revealing hybridisation events in Stipa (one of the largest genera in Poaceae). We demonstrate the usage of the classical morphological approach combining with scanning electron microscopy data, as well as analyses of pollen grains, and the application of molecular techniques including data derived from next generation sequencing. We illustrate the application of this integrative tool by specific examples: (1) in the S. × heptapotamica hybrid complex comprising morphologically distant but genetically closely related species, and (2) in the S. × lazkovii hybrid complex containing genetically distant species
Marcin Nobis
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Achnatherum на территории Алтайской горной страны. Установлено, что на данной территории род представлен 5 видами. Составлены карты распространения и оригинальный ключ для определе-ния видов рода на территории Алтайской горной страны. Ключевые слова. Алтайская горная страна, злаки, ключ, распространение, таксономия, Achnatherum. Summary. Here we present a taxonomic revision of the genus Achnatherum in the Altai Mountain Country. On this area, the genus is represented by five species. Distribution maps of the examined taxa and a key for their identification are also provided.
Marcin Nobis
added a research item
The genus Stipa L. comprises over 150 species, all native to the Old World, where they grow in warm temperate regions throughout Europe, Asia, and North Africa. It is one of the largest genera in the family Poaceae in Middle Asia, where one of its diversity hotspots is located. However, identification of Middle Asian Stipa species is difficult because of the lack of new, comprehensive taxonomic studies including all of the species recorded in the region. We present a critical review of the Mid-Asian representatives of Stipa, together with an identification key and taxonomic listing. We relied on both published and unpublished information for the taxa involved, many of which are poorly known. For each taxon, we present a taxonomic and nomenclatural overview, habitat preferences, distribution, altitudinal range, and additional notes as deemed appropriate. We describe four new nothospecies: S. ×balkanabatica M. Nobis & P. D. Gudkova, S. ×dzungarica M. Nobis, S. ×pseudomacroglossa M. Nobis, S. ×subdrobovii M. Nobis & A. Nowak, one subspecies S. caucasica Schmalh. subsp. nikolai M. Nobis, A. Nobis & A. Nowak, and eight varieties: S. araxensis Grossh. var. mikojanovica M. Nobis, S. caucasica var. fanica M. Nobis, P. D. Gudkova & A. Nowak, S. drobovii (Tzvelev) Czerep. var. jarmica M. Nobis, S. drobovii var. persicorum M. Nobis, S. glareosa P. A. Smirn. var. nemegetica M. Nobis, S. kirghisorum P. A. Smirn. var. balkhashensis M. Nobis & P. D. Gudkova, S. richteriana Kar. & Kir. var. hirtifolia M. Nobis & A. Nowak, and S. ×subdrobovii var. pubescens M. Nobis & A. Nowak. Additionally, 12 new combinations, Achnatherum haussknechtii (Boiss.) M. Nobis, A. mandavillei (Freitag) M. Nobis, A. parviflorum (Desf.) M. Nobis, Neotrinia chitralensis (Bor) M. Nobis, S. badachschanica Roshev. var. pamirica (Roshev.) M. Nobis, S. borysthenica Klokov ex Prokudin var. anomala (P. A. Smirn.) M. Nobis, S. holosericea Trin. var. transcaucasica (Grossh.) M. Nobis, S. kirghisorum P. A. Smirn. var. ikonnikovii (Tzvelev) M. Nobis, S. macroglossa P. A. Smirn. var. kazachstanica (Kotuchov) M. Nobis, S. macroglossa var. kungeica (Golosk.) M. Nobis, S. richteriana var. jagnobica (Ovcz. & Czukav.) M. Nobis & A. Nowak, and S. zalesskii Wilensky var. turcomanica (P. A. Smirn.) M. Nobis are proposed, and the lectotypes for 14 taxa (S. arabica Trin. & Rupr., S. bungeana Trin. ex Bunge, S. caspia K. Koch, S. ×consanguinea Trin. & Rupr., S. effusa Mez, S. ×heptapotamica Golosk., S. jacquemontii Jaub. & Spach., S. kungeica Golosk., S. margelanica P. A. Smirn., S. richteriana, S. rubentiformis P. A. Smirn., S. sareptana A. K. Becker, S. tibetica Mez, and Timouria saposhnikovii Roshev.) are designated. In Middle Asia the genus Stipa comprises 98 taxa, including 72 species, four subspecies, and 22 varieties. Of the 72 species of feather grasses, 23 are of hybrid origin (nothospecies). In Middle Asia, feather grasses can be found at elevations from (0 to)300 to 4500(to 5000) m, but most are montane species. The greatest species richness is observed at altitudes between 1000 and 2500 m. Nineteen species grow above 3000 m, but only nine above 4000 m. The number of taxa (species and subspecies) growing in each country also varies considerably, with the highest noted in Kazakhstan (42), Tajikistan (40), and Kyrgyzstan (35). Of the 76 taxa of Stipa (species and subspecies) recorded in Middle Asia, 41 are confined to the region, with some being known only from a single country or mountain range. Distribution maps of selected species are provided.
Marcin Nobis
added a research item
Increasing urbanisation has led to the fragmentation of natural habitats and biodiversity decline. In Central Europe, this fate has befallen semi-natural xerothermic grasslands. However, in the case of species well adapted to open habitats, urbanisation may positively affect dispersal. In many places specialised xerothermic species start to spread in secondary habitats such as roadsides or railway embankments. However, little is known about the genetic variability of such newly-established populations. In an era of growing anthropogenic pressure, understanding the role of secondary, anthropogenic habitats in the maintenance biodiversity should be considered a crucial issue in the conservation of species associated with Central European dry grasslands. Therefore, on the basis of a combined analysis of xerothermic species abundance, bioclimatic and soil data, and genetic diversity, we wish to examine the suitability of secondary habitats (as substitutes for primary) and to demonstrate their role in the conservation of xerothermic species. Our results suggest that smaller competition for light and greater habitat connectivity indicate suitability of habitats for xerothermic species conservation. On one hand, the occurrence of these species in secondary habitats may be limited by the pressure of expansive grasses, on the other, the negative impact of the latter is buffered by the reduced presence of shrubs, and thus the greater availability of light. Plant communities from secondary habitats are less richness in xerothermic species, and particular species are less abundant, for this reason secondary habitats should be treated more as refuges (with suboptimal conditions) than as substitution habitats. But yet, Stipa pennata populations from secondary habitats do not suffer from genetic depauperation. Populations from secondary habitats are characterised by comparable share of two genotypes, indicating existence of gene flow between populations from south and north-west part of studied area. Through the maintenance of meta-population processes by means of dynamic connectivity between habitats on a landscape scale, secondary habitats possess an undoubtedly high conservation value for xerothermic species. https://authors.elsevier.com/a/1aGTI,XRNLctLo
Marcin Nobis
added a research item
Stipa splendens (syn. Achnatherum splendens) has been transferred to Neotrinia (Tzvelev) M. Nobis, P. Gudkova et A. Nowak gen. nov. as N. splendens (Trin.) M. Nobis, P. Gudkova et A. Nowak, based on a comparison of the macromorphology and lemma micromorphology of the Asian representatives of the tribe Stipeae. Lemma epidermal patterns in the examined species are presented and discussed. Additionally, a new section Pennatherum M. Nobis comprising Achnatherum pelliotii, a species recently transferred to Achnatherum from Ptilagrostis is also proposed. This taxon clearly differs from Ptilagrostis by the lemma micromorphology. The species with typical maize-like lemma micromorphological pattern is apparent member of Achnatherum, however, because of having plumose awns and short, blind calluses, it was earlier included into Ptilagrostis. Achnatherum pelliotii is the only Old Word Achnatherum species, with long pilose awns, thus, we propose to place it in the new section. Neotrinia gen. nov. и Pennatherum sect. nov. рода Achnatherum (Poaceae: Stipeae) Ключевые слова: новая секция, новый род, таксономия, Poaceae, Stipeae. Аннотация. В статье приводится новый род Neotrinia (Tzvelev) M. Nobis, P. Gudkova et A. Nowak gen. nov., принадлежащий трибе Stipeae (Poaceae), включающий N. splendens (Trin.) M. Nobis, P. Gudkova et A. Nowak, (syn. Stipa splendens, Achnatherum splendens). Описание основано на сравнении макро-и микроморфологиче-ского строения нижних цветковых чешуй азиатских представителей трибы Stipeae. Также представлено об-суждение паттернов микроморфологического строения нижних цветковых чешуй исследованных (азиатских) видов трибы Stipeae. Кроме того, описана новая секция Pennatherum M. Nobis рода Achnatherum, включающая A. pelliotii, вид, недавно переведенный в Achnatherum из близкородственного рода Ptilagrostis. Achnatherum pelliotii отличается от Ptilagrostis по микроморфологическому строению нижних цветковых чешуй и относит
Ewelina Klichowska
added a research item
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.
Marcin Nobis
added a research item
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.
Jakub Sawicki
added a research item
The article takes up the problem of deficiency of molecular marker, which could illustrate molecular variability as well as phylogenetic relation within the genus of Stipa L. (Poaceae). Researches made so far hadn’t delivered sufficient information about relationships between particular taxa from the genus of Stipa. In the present study, we analyzed variability and phylogenetic informativeness of nuclear ribosomal DNA in six species from the genus against five other species from Poaceae including a division of this region into functional elements and domains. Our results showed that the intergenic spacer region, and especially its part adjacent to 26 S nrDNA, is a molecular marker giving a real chance for a phylogeny reconstruction of Stipa. The region seems to be the most phylogenetically informative for Stipa from all the chloroplast and nuclear markers tested so far. Comparative analysis of nrDNA repeat units from Stipa to other representatives of Poaceae showed that their structure does not deviate from the general scheme. However, the rate of evolution within the inter-repeats in the IGS region is extremely high and therefore it predestines the region for phylogenetic analyses of Stipa at genus level or in shallower taxonomic scale.
Ewelina Klichowska
added a research item
Citation: Klichowska E, Nobis M (2017) Stipa pennata subsp. ceynowae (Poaceae, Pooideae), a new taxon from Central Europe. PhytoKeys 83: 75–92. https://doi. Abstract 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. pen-nata 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.
Marcin Nobis
added 3 research items
Stipa klimesii sp. nov. from the Western Himalayas (India: Ladakh) and its variety S. klimesii var. pubescens var. nov. are described. The new species is similar to S. roborowskyi but differs by its longer anthecium, longer ligules of vegetative shoots, longer hairs on seta and shorter awns. Stipa klimesii is also similar to S. purpurea, but differs by longer ligules of vegetative shoots, shorter awns, slightly shorter hairs on seta and by the character of the panicle, which is compressed and with straight branches in S. klimesii vs. lax and with flexuous branches in S. purpurea. Epidermal patterns of the lemma in S. klimesii and both above-mentioned species, were examined by scanning electron microscopy. Images of macromorphological and micromorphological structures are provided.
Stipa ×fallax nothosp. nov. (Poaceae), from western Pamir Alai Mts (Tajikistan), is described and illustrated. Field observation, numerical analyses of morphology, and pollen grain viability data show that it originated from hybridization between S. drobovii and S. macroglossa subsp. macroglossa, species representing sections Smirnovia and Stipa, respectively. Stipa ×fallax is morphologically close to S. ×alaica and S. ×hissarica, but is distinguished by its shortly pilose lower part of the awn and densely pubescent leaves. Characters distinguishing S. ×fallax from its parental species as well as similar hybrid taxa in section Smirnovia that also grow in Central Asia are presented. The micromorphology of lemmas and leaves of S. ×fallax and its parental species was examined by scanning electron microscopy. We also propose the new combination Stipa drobovii var. iskanderkulica (Tzvelev) M.Nobis & A.Nowak.
Marcin Nobis
added a research item
Stipa krylovii is newly reported for the flora of Nepal, and this is the most southerly location yet found for this species. A full description of S. krylovii is included, along with illustrations, notes on its taxonomy and a distribution map.
Marcin Nobis
added a research item
The genus Stipa comprises approximately 50 taxa in Kazakhstan, of which 18 were described by Yuri A. Kotukhov from eastern Kazakhstan between 1987 and 1998. The majority of the original material of these taxa is preserved in the LE herbarium (Saint-Petersburg, Russia). Some original material of Kotukhov’s species, not mentioned in the protologues, is preserved in the KUZ (Kemerovo, Russia) and KRA (Krakow, Poland) herbaria. The present paper presents a complete list of feather grasses described by Kotukhov from Kazakhstan, with indications of types and the locations where they are preserved, lectotypification of seven names (S. akseirica, S. argillosa, S. karakabinica, S. kempirica, S. kyzylkiensis, S. azutavica, S. saikanica), one new combination (S. orientalis var. azutavica comb. nova), synonymisation of three species (S. akseirica is a synonym of S. sareptana, S. kyzylkiensis is a synonym of S. sczerbakovii, and S. saikanica is a synonym of S. lessingiana), an identification key and taxonomic notes regarding the discussed taxa.
Marcin Nobis
added 9 research items
The Stipa section Smirnovia Tzvel. includes 32 taxa. Of this number, 18 taxa, with glabrous (smooth or scabrous) awn columns, belong to the Stipa lipskyi group, which is the object of the present study. The centre of diversity of the studied group is located in the mountains of Middle Asia (Pamir Alai and Tian-Shan). Based on the examination of a large number of herbarium specimens (over 550 sheets), field observations, scanning electron microscopy observation of lemma micromorphology and statistical analysis of macromorphological characters, 12 taxa at the specific level were recognized on the area studied. Three species of hybrid origin, Stipa × tadzhikistanica M. Nobis, S. × brevicallosa M. Nobis, and S. × hissarica M. Nobis, as well as the variety S. lipskyi var. pilosivaginata M. Nobis, were described as new to science. Additionally, lectotypes for S. karataviensis Roshev. and S. longiplumosa Roshev. were designated and a new combination Stipa macroglossa subsp. kazachstanica (Kotuch.) M. Nobis was proposed. Numerous new localities, often far from the previously known distribution area, were reported for several species. Illustrations and a key to species and varieties are provided in addition to complete descriptions including information on nomenclatural types, synonymies, distributions, habitat and specimens examined.
Morphological investigations were conducted on four taxa of the Stipa tianschanica complex occurring in Central Asia. The main objective of the study was to survey the micromorphological characters of the lemma in the complex by means of scanning electron microscopy (SEM) in order to assess their taxonomic value and to compare the variation with the recent treatment based on macromorphological traits. The main lemma epidermal characters studied included the frequency, shape and distribution of long cells, hooks, prickles and macrohairs. A principal component analysis of macromorphological traits revealed correlations with micromorphological characters and with the geographical distribution of the specimens examined. On this basis, three species, namely S. tianschanica, S. klemenzii and S. austromongolica, and one additional subspecies, S. tianschanica subsp. gobica, are recognized in Central Asia. Stipa austromongolica is described here as a species new to science. The lectotype for S. tianschanica is designated here as well. Morphological descriptions, synonyms, distribution maps, citations of representative specimens and an identification key are provided.
Stipa ×brozhiana M. Nobis nothosp. nov. (Poaceae) is described and illustrated. The taxon belongs to sect. Smirnovia Tzvel. and originated from the hybridization of species belonging to sect. Smirnovia and Barbatae Junge. Stipa ×brozhiana is morphologically close to Stipa lipskyi Roshev., but is easily distinguished by its much shorter hairs on the seta, thinner, uni- or indistinctly bigeniculate awns, differently shaped callus and longer ligules of vegetative shoots. Characters distinguishing S. ×brozhiana from the parental species and other similar hybrid taxa belonging to sect. Smirnovia and occurring in the Pamir Alai Mts are discussed. In addition, the taxonomical status of S. ×tzvelevii Ikonn. pro sp., another taxon that has originated from hybridization between species belonging to sect. Smirnovia and Barbatae, is discussed. The taxon was originally described as a distinct species, but is now considered to be the hybrid S. caucasica×S. orientalis. The main characters distinguishing S. ×tzvelevii from parental species are given.
Marcin Nobis
added 2 research items
Numerical analyses of macromorphological characters of specimens of S. lessingiana, S. borysthenica and S. baktashevae, nested the last one among individuals of S. borysthenica. We propose synonymization of S. baktashevae, the species described in 2014 and known only from type material, with S. borysthenica, and provide morphological comparisons of the taxa.
Stipa dickorei sp. nov. from the Western Tibetan Plateau (China) is described. The new species is morphologically similar to S. regeliana, but they differ from each other in the length of ligules of vegetative shoots. Stipa dickorei is also similar to S. aliena, however they differ in the shape of panicle, which is contracted with straight branches in S. dickorei, and lax with flexuous branches in S. aliena. Images of macromorphological and micromorphological structures of the new taxon are provided. Additionally, new records of S. borysthenica, S. richteriana, and S. zalesskii, species not listed in the recent Flora of China, as well as a checklist of Chinese feather grasses are also presented.