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Flowers in three species of Aquilegia. A. A. kansuensis (photographed by B. Liu). B. A. yangii (photographed by B. Liu). C. A. oxysepala (photographed by D. Yakubov). Insets: flowers with the sepals and petals removed.

Flowers in three species of Aquilegia. A. A. kansuensis (photographed by B. Liu). B. A. yangii (photographed by B. Liu). C. A. oxysepala (photographed by D. Yakubov). Insets: flowers with the sepals and petals removed.

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Aquilegia yangii (Ranunculaceae), a new species from western China, is described and illustrated. The new species is similar to A. oxysepala and A. kansuensis. However, there are differences in reproductive structure, especially flower size, petal shape, anther colour and seed surface, between the new species and the other two species. The seed sur...

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... differs by having smaller flowers, yellow petal laminae (vs. white), yellow anthers and a seed coat irregularly swollen without obvious outlines of epidermis cells (Fig. 5). Stamens of A. oxysepala and A. kansuensis become dark in older flowers as anthers dehisce, whereas A. yangii is characterized by yellow anthers throughout its flowering time (Fig. 6). Individual plants or hybridogenetic populations characterized by intermediate features may occur in sites where the species' habitats are in contact. In rare cases, one of the parent species may be absent from such locations, due to the activity of insect pollinators. The morphological differences between the new species, A. oxysepala ...

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... For example, a recent study showed that the diversity of Asian Aquilegia is much higher than that previously recognized in different floristic research (Bulavkina 1937, Fu & Robinson 2001, Erst et al. 2017. In recent years, several new species have been described from China (A. hebeica Erst (2017: 127), A. xinjiangensis Erst (2017: 122), A. yangii Luo & Li (2018: 2018), India and Pakistan (A. bashahrica Erst (2020: 112), A. × emodi Erst (2020: 113)), Mongolia (A. daingolica Erst & Shaulo (2013: 15), A. grubovii Erst et al. (2016: 114)), and Russia (Aquilegia aradanica Shaulo & Erst (2011: 29), A. barykinae Erst et al. (2014: 110), A. kamelinii Erst et al. (2013: 20), A. synakensis Shaulo & Erst (2010: 43). Moreover, two new combinations, A. kansuensis (Brühl) Erst (2014: 25) and A. semicalcarata (Schipczinsky) Erst (2015: 112), have been proposed. ...
... For example, a recent study showed that the diversity of Asian Aquilegia is much higher than that previously recognized in different floristic research (Bulavkina 1937, Fu & Robinson 2001, Erst et al. 2017. In recent years, several new species have been described from China (A. hebeica Erst (2017: 127), A. xinjiangensis Erst (2017: 122), A. yangii Luo & Li (2018: 2018), India and Pakistan (A. bashahrica Erst (2020: 112), A. × emodi Erst (2020: 113)), Mongolia (A. daingolica Erst & Shaulo (2013: 15), A. grubovii Erst et al. (2016: 114)), and Russia (Aquilegia aradanica Shaulo & Erst (2011: 29), A. barykinae Erst et al. (2014: 110), A. kamelinii Erst et al. (2013: 20), A. synakensis Shaulo & Erst (2010: 43). Moreover, two new combinations, A. kansuensis (Brühl) Erst (2014: 25) and A. semicalcarata (Schipczinsky) Erst (2015: 112), have been proposed. ...
Article
We report the first record of Aquilegia grubovii (Ranunculaceae) for Russia, a rare species previously known from five localities in Mongolia. The species belongs to A. sect. Aquilegia subsect. Aquilegia and is characterized by curved or hooked spurs slightly longer than the limb of petals, 2-2.8 mm long, straight only at the top diverging fruits with stylodia 2 times shorter than follicles, smooth or tuberculate seed surface and stems covered with simple and glandular hairs. The new records of Aquilegia grubovii are located in the Azas Nature Reserve, which is situated in the central part of the Todzha basin, Tuva Republic (a large intermountain trough within the Altai-Sayan mountain region) of south-central Asia about 500 km west of Lake Baikal. We provide an assessment of the conservation status, geographical distribution and habitat of Aquilegia grubovii, along with an identification key to all species from Russia.
... In East Asia, Aquilegia has a monophyletic clade (including A. ecalcarata Maxim, A. yabeana Kitag., A. rockii Munz, A. kansuensis [Brühl] Erst, and A. yangii Luo and Li [Luo et al., 2018]) ( Figure 1) that is widely distributed in three subkingdoms of the East Asian flora. Aquilegia yabeana is mainly distributed in subkingdom iii (the Sino-Japanese Forest subkingdom), with the largest flowers, yellow dehiscent anthers, and spurs that curve into a hook at the apex (Figures 1 and 2a,b). ...
... Aquilegia yangii is mainly distributed in subkingdom i, with yellow petal blades, smaller flowers, and yellow dehiscent anthers. Aquilegia ecalcarata is the most widely distributed, and it is distributed in subkingdoms i, ii, and iii, with black dehiscent anthers and spurs that are absent or short ( Figures 1 and 2a,b) (Fu & Robinson, 2001;Luo et al., 2018;Wu et al., 2010;Wu & Wu, 1996;Xue et al., 2020). According to previous studies, A. ecalcarata is not a uniform group in terms of its genetics or floral traits Xue et al., 2020). ...
... According to previous studies, A. ecalcarata is not a uniform group in terms of its genetics or floral traits Xue et al., 2020). Based on 10 nuclear gene fragments, samples of A. ecalcarata were divided into two clades (western and eastern) that had the closest genetic relationship with A. rockii and A. kansuensis (some populations distributed in Gansu and Qinghai Provinces have been identified as a new species, A. yangii) Luo et al., 2018). Even in the west, the similarity of floral morphology between A. ecalcarata and A. rockii is greater than that of A. ecalcarata itself (western vs. eastern) (Xue et al., 2020). ...
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Quaternary climate oscillations and geographical heterogeneity play important roles in determining species and genetic diversity distribution patterns, but how these factors affect the migration and differentiation of East Asian plants species at the population level remains poorly understood. The Aquilegia ecalcarata complex, a group that originated in the Late Tertiary and is widely distributed throughout East Asia, displays high genetic variation that is suitable for studying elaborate phylogeographic patterns and demographic history related to the impact of Quaternary climate and geography. We used plastid genome data from 322 individuals in 60 populations of the A. ecalcarata complex to thoroughly explore the impact of Quaternary climate oscillations and geography on the phylogeographic patterns and demographic history of the A. ecalcarata complex through a series of phylogenetic, divergence time estimation, and demographic history analyses. The dry, cold climate and frequent climate oscillations that occurred during the early Pleistocene and the Mid‐Pleistocene transition led to the differentiation of the A. ecalcarata complex, which was isolated in various areas. Geographically, the A. ecalcarata complex can be divided into Eastern and Western Clades and five subclades, which conform to the divergence of the East Asian flora. Our results clearly show the impact of Quaternary climate and geography on evolutionary history at the population level. These findings promote the understanding of the relationship between plant genetic differentiation and climate and geographical factors of East Asia at the population level.
... Article ecalcarata is not at the base of the genus; instead, A. ecalcarata, A. yabeana, A. kansuensis and A. rockii comprise a clade (Fior et al., 2013). Luo et al. (2018) referred to A. kansuensis from Gansu and Qinghai provinces as A. yangii. A. ecalcarata has numerous overlaps in distribution with its four related species ( Supplementary Fig. 1), and the differences in floral phenotype between A. ecalcarata and its four related species are the size of the floral organs and the petal with or without nectar spurs (Fig. 1b). ...
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Traits are basic attributes of organisms that form the basis for speciation and diversity. The floral nectar spur is a classic example of a key innovative trait. Differences in nectar spur morphology can lead to pollinator specialization and thereby promote reproductive isolation between species. Despite its importance, the nectar spur has been lost in some members of the columbine genus (Aquilegia), resulting in a new spurless trait, and the evolutionary influence of this trait has become a topic of scientific interest. Aquilegia ecalcarata is an important representative columbine species that lacks spurs. Here, we resequenced the genomes of 324 individuals from A. ecalcarata and four related species. We found that A. ecalcarata was divided into three groups based on the phylogenetic relationships and population genetic structures. Topology weighting analysis revealed that A. ecalcarata has multiple origins, and Patterson's D statistic showed that the spurless trait may have one origin. Floral morphological analysis revealed significant differences between A. ecalcarata and its spurred sister groups, and the floral phenotypes of the three A. ecalcarata groups have identical or similar floral phenotypes. Our results confirmed that the spurless trait not only produced the phenotype of A. ecalcarata but also contributed to the emergence of the A. rockii phenotype. Moreover, the spurless trait promoted the divergence between A. ecalcarata and its close, spurred relatives. Our research shows that the loss of key innovative traits can play a very important part in speciation and species diversity. This article is protected by copyright. All rights reserved.
... The genus Aquilegia Linnaeus (1753: 533) (Ranunculaceae) consists of 110 species and is mainly distributed in the temperate regions of the Northern Hemisphere (Erst et al. 2020a), with 21 species in North America (Whittemore 1997), 34 in Asia (Erst et al. 2017, Luo et al. 2018, and 56 in Europe (Nardi 2016). According to the most recent taconomic treatment of Aquilegia (Erst et al. 2020a), there are six species and one hybrid of Aquilegia in Himalayas: A. moorcroftiana Wall. ...
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As part of the ongoing revision of the Himalayas species of Aquilegia (Ranunculaceae), three names applicable to three accepted species are typified, and some nomenclatural and taxonomic remarks are provided. Lectotypes are designated for A. glauca (a taxonomic synonym of A. fragrans) and A. vulgaris subsp. nivalis (a nomenclatural synonym of A. nivalis). Epitypes for A. fragrans and A. glauca were also designated in order to avoid any ambiguity in the taxonomic interpretation of the lectotypes and hence the application of the names.
... The genus Aquilegia L. (columbine), comprising approximately 70 perennial herb species, belongs to the family Ranunculaceae and is widely distributed in North America and Eurasia (Munz, 1946). Recently, several new species were reported, bringing the number of columbine taxa to about 110 species (Erst et al., 2017(Erst et al., , 2020Luo et al., 2018). Although the morphologies and habitats of columbine species differ, the phylogenetic resolution of this genus at the molecular level is very low, and therefore the genus is considered to be a widespread population complex. ...
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Premise: Aquilegia is an ideal taxon for studying the evolution of adaptive radiation. Current phylogenies of Aquilegia based on different molecular markers are inconsistent, and therefore a clear and accurate phylogeny remains uncertain. Analyzing the chloroplast genome, with its simple structure and low recombination rate, may help solve this problem. Methods: Next-generation sequencing data were generated or downloaded for Aquilegia species, enabling their chloroplast genomes to be assembled. The assemblies were used to estimate the genome characteristics and infer the phylogeny of Aquilegia. Results: In this study, chloroplast genome sequences were assembled for Aquilegia species distributed across Asia, North America, and Europe. Three of the genes analyzed (petG, rpl36, and atpB) were shown to be under positive selection and may be related to adaptation. The phylogenetic tree of Aquilegia showed that its member species formed two clades with high support, North American and European species, with the Asian species being paraphyletic; A. parviflora and A. amurensis clustered with the North American species, while the remaining Asian species were found in the European clade. In addition, A. oxysepala var. kansuensis should be considered as a separate species rather than a variety. Discussion: The complete chloroplast genomes of these Aquilegia species provide new insights into the reconstruction of the phylogeny of related species and contribute to the further study of this genus.
... kansuensis that was collected from Yuzhong County, Gansu Province, China, for whole-genome sequencing. We noticed that a new name, A. yangii, has recently been given to plants of this and many other populations 19 . However, we decided to stay with the original name because the new one is still poorly known. ...
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The genus Aquilegia (Ranunculaceae) has been cultivated as ornamental and medicinal plants for centuries. With petal spurs of strikingly diverse size and shape, Aquilegia has also been recognized as an excellent system for evolutionary studies. Pollinator‐mediated selection for longer spurs is believed to have shaped the evolution of this genus, especially the North American taxa. Recently, however, an opposite evolutionary trend was reported in an Asian lineage, where multiple origins of mini- or even nonspurred morphs have occurred. Interesting as it is, the lack of genomic resources has limited our ability to decipher the molecular and evolutionary mechanisms underlying spur reduction in this special lineage. Using long-read sequencing (PacBio Sequel), in combination with optical maps (BioNano DLS) and Hi–C, we assembled a high-quality reference genome of A. oxysepala var. kansuensis, a sister species to the nonspurred taxon. The final assembly is approximately 293.2 Mb, 94.6% (277.4 Mb) of which has been anchored to 7 pseudochromosomes. A total of 25,571 protein-coding genes were predicted, with 97.2% being functionally annotated. When comparing this genome with that of A. coerulea, we detected a large rearrangement between Chr1 and Chr4, which might have caused the Chr4 of A. oxysepala var. kansuensis to partly deviate from the “decaying” path that was taken before the split of Aquilegia and Semiaquilegia. This high-quality reference genome is fundamental to further investigations on the development and evolution of petal spurs and provides a strong foundation for the breeding of new horticultural Aquilegia cultivars.
... The genus Aquilegia Linnaeus (1753: 533) (Ranunculaceae) consists of about 110 species and is mainly distributed in the temperate regions of the Northern Hemisphere (Luo et al. 2018), with 21 species in North America (Whittemore 1997), 34 in Asia (Erst et al. 2017, Luo et al. 2018, and 56 in Europe (Nardi 2016). Aquilegia (known as columbines) possesses unusual floral organs, represented by petaloid sepals, staminodium, and petals with a nectar spur, which allow the study of evolutionary novelties (Kramer et al. 2007;Voelckel et al. 2010). ...
... The genus Aquilegia Linnaeus (1753: 533) (Ranunculaceae) consists of about 110 species and is mainly distributed in the temperate regions of the Northern Hemisphere (Luo et al. 2018), with 21 species in North America (Whittemore 1997), 34 in Asia (Erst et al. 2017, Luo et al. 2018, and 56 in Europe (Nardi 2016). Aquilegia (known as columbines) possesses unusual floral organs, represented by petaloid sepals, staminodium, and petals with a nectar spur, which allow the study of evolutionary novelties (Kramer et al. 2007;Voelckel et al. 2010). ...
... For example, a recent study showed that the diversity of Chinese Aquilegia is much higher than that previously recognized (Fu & Robinson 2001, Erst et al. 2017. This study revealed that instead of the 13 species previously recognized from China there are 22 species (Erst et al. 2017, Luo et al. 2018, even with the exclusion of two species previously thought to occur China (Erst et al. 2017). It is therefore likely that many endemic species remain to be found in mountainous areas of Asia, ...
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A new species Aquilegia bashahrica and a new nothospecies Aquilegia × emodi from North-western Himalayas, are described and illustrated. In addition A. lactiflora is recorded for India and Pakistan for the first time and Aquilegia kareliniana is excluded from the flora of India. An identification key to the species of Aquilegia from the North-western Himalayas is provided and diagnostic characters are discussed.
Article
Aquilegia ecalcarata Maxim. is the only spurless species within the genus Aquilegia and comprises a monophyletic clade with A. yabeana Kitag., A. kansuensis Brühl, and A. rockii Munz. Our previous study on the genetic diversity of those four species revealed that the populations of A. ecalcarata can be divided into two groups, indicating possible genetic difference within A. ecalcarata. However, it is not clear whether the genetic difference is related to the morphological variation among species and groups of A. ecalcarata populations. To answer that question, the morphological variation patterns based on 22 floral and 19 vegetative traits from 42 populations, covering the entire distribution of A. ecalcarata and its relatives, were analyzed in the present study. The result showed that: (i) the differences among the four species were reflected in the floral rather than the vegetative traits; (ii) populations of A. yabeana and A. rockii fell into one cluster each, and each of the six clusters occupied its own distribution range; (iii) one of two A. ecalcarata clusters fell into a subgroup and shared common floral traits with A. rockii; (iv) the individuals of A. ecalcarata form. ecalcarata and form. semicalcarata were often mixed in the same population; and (v) the populations of A. kansuensis were split into two clusters, which differed obviously in floral traits. These results will provide an important morphological basis for the redefinition of species and lay a foundation for the further exploration of the “spurless” A. ecalcarata.