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Article

Endemic of Kazakhstan Allium lehmannianum Merckl. Ex Bunge and Its Position Within the Genus Allium

by
Daulet Sh. Abdildanov
1,2,
Polina V. Vesselova
1,
Gulmira M. Kudabayeva
1,
Bektemir B. Osmonali
1,
Michail V. Skaptsov
3 and
Nikolai Friesen
4,*
1
Institute of Botany and Phytointroduction, 36D Timiryazev str., Almaty 050040, Kazakhstan
2
Department of Biodiversity and Bioresources, Faculty of Biology and Biotechnology, Al-Farabi Kazakh National University, Almaty 050040, Kazakhstan
3
South Siberian Botanical Garden, Faculty of Biology, Altai State University, Barnaul 656906, Russia
4
Botanical Garden, School of Biology/Chemistry, Osnabruck University, 49076 Osnabruck, Germany
*
Author to whom correspondence should be addressed.
Plants 2025, 14(7), 1113; https://doi.org/10.3390/plants14071113
Submission received: 19 February 2025 / Revised: 19 March 2025 / Accepted: 1 April 2025 / Published: 2 April 2025
(This article belongs to the Special Issue Plant Diversity and Classification)
Browse Figures
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Abstract

:
The paper presents data on the phylogeny, anatomy of vegetative organs, phytocenosis composition, and ploidy of two populations of endemic Allium lehmannianum Merckl. ex-Bunge. in the flora of northeastern Betpakdala. Some discrepancies in nomenclature and sectional affiliation are discussed and corrected. Allium lehmannianum belongs to the section Multicaulea F. O. Khass. and & Yenglal. subgenus Allium, the main range covering the eastern Old Mediterranean region. The ploidy of Allium lehmannianum was studied for the first time using flow cytometry techniques. The sequencing of nrITS was used to construct a phylogenetic tree, including sequences from the NCBI database. The phylogenetic tree of A. lehmannianum was compiled by taking into account previously published data. During the fieldwork in the northeastern part of Betpakdala, the species was found to grow in a rubbly, stony, weakly undulating plain in two different communities. In this work, we applied molecular genetic, cytometric, and anatomical studies on the collected samples of two populations from the northeastern part of the Betpakdala desert to determine the taxonomic composition of a relict and endemic species of Kazakhstan.

1. Introduction

The uniqueness of the flora of a particular region is based on the set of endemic species that are suited to the growing conditions. In the flora of Kazakhstan, 130 onion species have been noted [1], 21 of which are endemic to the territory of the Republic [2]. At the same time, most onion species, both in general and directly endemic, are confined to mountainous areas [3]. In the desert part of Kazakhstan, three endemic species grow—Allium lehmannianum Merckl. ex Bunge (section Multicaulea F. O. Khass. and & Yenglal.), A. iliense Regel (section Regeloprason Wendelbo), A. subscabrum (Regel) R.M.Fritsch (section Desipienta (Omelchuk), and R.M.Fritsch subsection Viceniprason R.M.Fritsch and N.Friesen) [4,5]. Moreover, if the last two species belong to the group of psammophytes, then A. lehmannianum prefers petrophytic and clayey habitats, usually found on rubble-rocky substrates.
Here, it is necessary to note some discrepancies in the sectional assignment of Allium lehmannianum and the nomenclature of the sections Multicaulea and Unicaulea F.O.Khass. Initially, F. Khassanov, when describing the section Multicaulea F. O. Khass. and & Yenglal. [6], cited Allium lehmannianum as the type of species of the section. In 2011, Khassanov and his colleagues [7] described the new species A. ravenii and the new section Unicaulea with the type of species A. kotschyi Boiss. Aftewards Khassanov et al. [8,9] indicated that Allium lehmannianum belongs to the section Unicaulea, subgenus Allium, together with other species such as A. kotschyi Boiss and a new species for the flora of Uzbekistan, A. ravenii F. O. Khass., Shomur. et Kadyrov. Allium ravenii is morphologically close to the studied species, distinguished by a smaller flower size and the absence of bracts [7]. In the new flora of Uzbekistan [10], for the section Multicaulea, a new type of species, A. borszczowii, is given for the first time without any explanation, which is a violation of the Code of Nomenclature. The type of species is not arbitrarily changed [11]. Thus, A. lehmannianum belongs to the section Multicaulea, and the section Unicaulea should probably be a synonym of the section Multicaulea, since the type of species A. kotschyi, judging by the herbarium specimens viewed in GBIF https://www.gbif.org/species/2856108 (accessed on 10 January 2025) [12], is morphologically very close to A. lehmannianum and A. ravenii. In the latest work by Khassanov et al. [9], which is dedicated to both sections, the type of species for the section Unicaulea is already given as A. lehmannianum instead of A. kotschyi legitimately given in the section description. As for the geographical confinement of A. lehmannianum, its range is indicated differently in different sources. For example, Vvedensky in the “Flora of the USSR” [13] lists Allium lehmannianum (section Porrum Don.) for the Kyzylkum and Aral-Caspian (the species is described from the Aral deserts: Lectotype (LE 00053884): “Lehmfläche zwischen dem Jaxarts und dem See Aigiräk, 15 May 1842”, which was isolated from three sheets stored in the LE Herbarium in Sankt-Peterburg. Thus, according to Khassanov et al. [9], the information about the locus classicus of the studied species is as follows: “Clay surface between Jaxartes and Lake Aigirak Aral deserts”, which was written on the label of A. Bunge, and they in 1840, went to the eastern coast of the Caspian Sea, where they constantly carried out various excursions and collected rich materials and collections of fauna and flora in the Northern Aral Sea region and Mangyshlak [9]. Also, according to the expedition route and local names, Lake Aigyrak could have been located on the site of a swamp, which on maps from the mid-19th century is designated as a lake in the lower reaches of the Syr Darya River (Kazakhstan).
In the Identifier of Central Asian Plants [14], where the genus Allium was also processed by Vvedensky, the following region is already indicated: “Betpakdala, Priaralskie desert”. For the “Flora of Kazakhstan” [15], onions were processed by Pavlov and Polyakov, and in the column on the distribution of A. lehmannianum (section Porrum Don.), they indicate: Turgai, Western small hills, Kyzylorda, Betpakdala, and Kyzylkum floristic regions. At present, Khassanov et al. [9] indicated Kyzylkum, Syr Darya Karatau, Betpakdala, and Balkhash regions; however, there are no herbarium specimens in Kyzylkum, Kyzylorda region, and small western hills. Thus, it is reliably known (confirmed by herbarium specimens) (Appendix A) that Allium lehmannianum grows in the following regions: Turgai, Betpakdala, Shu-Ili, and Karatau (Figure 1). Interestingly, the listed territories are ancient geological formations that have undergone peneplainization [16]. The studied species is usually confined to rubble-rocky habitats and is quite rare, and despite the fairly wide representation of such habitats, it should also be noted that the height of desert hills limits the altitudinal range of the species. Combining the considered ecological and geographical characteristics allows us to classify this species as a relict element of the flora and designate its range as the eastern ancient Mediterranean [17,18]. During fieldwork in 2024 in the northeastern part of Betpakdala, this species was found in a rubble-rocky habitat.
Our work aimed to clarify the nomenclature, determine the position of the A. lehmannianum in the genus Allium system, and describe its anatomical and cytometric characteristics using molecular genetic, cytometric, and anatomical methods.

2. Results

2.1. Association of Allium lehmannianum with Plant Communities

During fieldwork in the northeastern part of the Betpakdala desert, as already noted, two populations with A. lehmannianum were discovered (Appendix B). In the first of them (N 46.000268 E 72.792945), this species grew in a petrophysics-perennial saltwort-wormwood community dominated by Artemisia terrae-alba Krasch. The community’s total projective cover (TPC) was 10–15%. The total number of species participating in forming this community is 18 (Appendix C), among which representatives of the Amaranthaceae family predominated. In the second population (N 46.600332, E 74.414978), A. lehmannianum was part of the stipaetum–krascheninnikoviosum community of a gently undulating plain with rare rocky hills with the dominance of Stipa caucasica Schmalh. The total number of species was also 18, most belonging to the Amaranthaceae family. The TPP of this community was 50–55%. Analyzing the species composition of these communities, it should be noted that they have common species: Artemisia terrae-albae, Astragalus campylotrichus Bunge, Bassia prostrata (L.) Beck, Oreosalsola arbusculiformis (Drobow) Sennikov, and Tulipa biflora Pall. (Figure 1).
In total, 30 species from 16 families were identified in the communities under consideration. The percentage ratio of the number of species of the leading families is shown in Figure 2. The quantitative predominance of the representatives of the Amaranthaceae family over the following three groups, moreover, more than twice, reflects the characteristic feature of the composition of the desert flora of Kazakhstan.

2.2. Anatomical Analysis

The cross-section of the A. lehmannianum leaf (Figure 3A) is rounded. A fairly dense cuticle layer indicates the xerophytic nature of the species. The epidermal cells (E) located in a single row under it are large in size (Table 1). Perpendicular to the epidermal cells in the centripetal direction located tightly adjacent to each other are long cells of the columnar mesophyll (PM). They are followed by small and large vascular bundles (Figure 3B) and then the main parenchyma (PX).
The stem of A. lehmannianum (Figure 3C) also has a rounded shape in cross-section. It should be noted that, like the leaf, the anatomical section of the stem has a dense cuticle layer. However, its spongy mesophyll (SM) cells are small (Table 2), and the outer vascular bundles and sclerenchyma cells (SC) are located around the entire circumference of the stem. At the same time, the inner vascular bundles (Figure 3D) are larger than the outer ones. The vascular bundles of the leaf and stem have a closed-collateral structure.
When studying the anatomical structure of the leaf and stem of A. lehmannianum, biometric measurements of the cells and their layers of two populations were performed (Table 1 and Table 2). Thus, when comparatively analyzing the biometric indicators of the leaf and stem of the studied species, it was revealed that the size of the cells of the leaf blade in the second population is larger than in the first population. In the biometric data of the stem, minor differences are observed in the two populations.

2.3. Flow Cytometry

The DNA content was determined in two populations of A. lehmannianum. The results are presented in Table 3.
The results obtained via flow cytometry showed the presence of diploids in two populations (Table 3), with an average DNA content of 35,323 pg in A. lehmannianum. In addition, one sample from population 2 is a probable triploid with a DNA content of 50,212 pg. Examples of uncompressed histograms of the studied A. lehmannianum samples are shown in Figure 4.

2.4. Phylogenetic Analysis

We sequenced nrITS by using five plants of A. lehmannianum from two populations and performed the phylogenetic analysis with all ITS sequences of the subgenus Allium available in the NCBI GenBank. A total of 111 sequences from the subgenus Allium and 6 sequences were selected as an outgroup (A. ramosum L., A. tuberosum L., and A. trifurcatum (F.T.Wang and Tang), J.M.Xu, A. oreoprasum Schrenk from the subgenus Butomissa, and A. rubens Schrad and A. vodopjanovae N.Friesen from the subgenus Rhizirideum). The alignment length of 117 ITS sequences is 691 characters; 213 characters are constant, 89 are parsimony-informative, and 389 are parsimony-informative. Unweighted parsimony analysis resulted in 2592 most parsimonious trees of 2678 steps (consistency index CI = 0.35; retention index RI = 0.79). AIC chose the substitution model HKY+G in jModelTest2 for the Bayesian analysis. The resulting phylogenetic tree (Figure 5) divides the subgenus Allium into three monophyletic groups, where the closest to the outgroup is the smallest clade with species of the sections Coerulea (Omelczuk) F.O.Khass., Haneltii F.O.Khass., and Eremoprasum (Kamelin) F.O.Khass., R.M.Fritsch, and & N.Friesen. Allium lehmannianum is placed in the second group, with the most significant section Allium. Basal is a small clade of species from the section Avulsea F.O.Khass., followed by species from the section Allium and species from the sections Multicaulea, Crystallina F.O.Khass. and Yengal., and Brevidentia F.O.Khass. and Yengal., which form two sister clades. All five accessions of A. lehmannianum (sect. Multicaulea) are very well separated from the neighbouring sections of Brevidentia and Crystallina. Only A. borszszowii (sect. Multicaulea after [10]) stands separately in this clade. The third group is not very strongly supported (0.97 PP and 87 BS) and consists of the sections Mediasia F.O.Khass., R.M.Fritsch and N.Friesen, Brevispatha Valsecchi, Pallasia (Tzag.) F.O.Khass., R.M.Fritsch and & N.Friesen, Scorodon K.Koch, Koppetdagia F.O.Khass., and partly of species from the sections Coerulea and Avulsea and from the well-supported (1.0 PP and 100 BS) species-rich section Codonoprasum Rchb.

2.5. CP Analysis

We sequenced two plastid regions by using A. lehmannianum: trnL gene, partial sequence, and the trnL-trnF intergenic spacer (trnL) and psbA-trnH spacer (see the NCBI GenBank accessions number in Table A2). There are few published sequences of these CP regions from the subgenus Allium, so we could only analyze the similarity of the sequenced CP regions using nucleotide blast analysis (https://blast.ncbi.nlm.nih.gov, accessed on 3 January 2025). Blast analysis of the CP trnL region from A. lehmannianum shows a very high similarity with A. filidens Regel from the sections Chrystallina (99.37%), A. nikolai F.O.Khass. (99.21%), and A. michaelis F.O.Khass. and Tojibaev from the section Brevidentia (99.28%), A. ferganicum Vved. from the section Multicaulea (99.28%), and A. sabulosum Steven ex Bunge from the section Eremoprasa (99.27%). Blast analysis of the CP region psbA-trnH from A. lehmannianum shows 100% identity with A. nikolai from the section Brevidentia and A. filidens from the section Crystallina and with many Allium species from the third evolutionary lineage with a higher than 99.5% similarity.

3. Discussion

There are not many anatomical studies of species of the subgenus Allium, so we cannot say much about the peculiarities of the anatomical data of A. lehmannianum. However, some studies [19,20] show the similarity of the anatomical structure of vegetative organs (leaf and stem) of A. lehmannianum for species of the subgenus Allium. There is a comparative difference of A. lehmannianum from other studies [19,20,21,22,23] in the biometric indices of leaf cells and stem anatomical structure. Examples are that in the leaf lamina’s transverse section, the vascular bundles are arranged in a circle (Figure 3A). In contrast, in the anatomical structure of the stem, the vascular bundles are arranged in two rings (Figure 3C); the outer conducting bundles are smaller than the inner one (Table 2). We attribute this to the ecological growth of the studied species.
Also, according to the database (https://cvalues.science.kew.org, accessed on 5 January 2025), in the genus Allium, the DNA amounts vary from a minimum of 15.20 (pg) to a maximum of 148.90 (pg). The average DNA amount in onions is 40.66 (pg). The DNA abundance of the subgenus Allium is poorly understood, but there are data on the DNA abundance of some species from this subgenus. For example, the amount of DNA in Allium caeruleum is 23.50 pg [22], while in Allium caesium, it is up to 25.90 pg [24,25]. Also, Allium sativum has 32.50 pg [22] and 49.40 pg for Allium filidens [26]. Our study of the average DNA content of A. lehmannianum in the first population was 35.323 pg and agrees with other Allium subgenus species. However, one sample from population 2 represents a probable triploid, so the DNA content is 50,212 pg (Table A2). Careful re-analysis of all species based on extensive material will be necessary to clarify the taxonomic value of genome size data in the subgenus Allium. The triploid plant found in the second population of A. lehmannianum indicates possible hybridization or disruption in meiosis. This population should be examined much more closely to determine the cause. Despite the finding that closely related species in the genus Allium can exhibit substantial differences in their DNA content, e.g., A. coeruleum and A. caesium [22,24], the genome size provides important data for taxonomic and phylogenetic analysis.
Molecular data (ITS and plastids) clearly show that A. lehmannianum is closely related to species from the sections Brevidentia, Crystallina, and Eremoprasa, and despite being a strongly under-protected clade Allium borszszowii, which belongs to the section Multicaulea [17], it is somewhat separate from all species from the sections Brevidentia, Chrystallina, and A. lehmannianum (Sect. Multicaulea). Together with other non-monophyletic sections (Caerulea, Avulsea) (Figure 5), the classification in the subgenus Allium still needs a lot of corrections. In addition, according to Khassanov [20], more than 70% of species from the subgenus Allium (375 species in the subgenus Allium), that have not yet been molecularly examined should be included in the analysis. Unfortunately, we could not examine other species (A. kotschyi, A. ravinii, A. ferganicum, A. rinae, A. oxianum) from the section Multicaulea. In our estimation, the sequencing of most species of the subgenus Allium (here, actually, the sequencing of ITS and some chloroplast fragments is sufficient) will reveal many taxonomic and phylogenetic surprises inside the subgenus Allium.

Nomenclatural Notice

Allium sect. Multicaulea F.O. Khass. and Yengal., 1996 in M.A. Öztürk, Ö. Seçmen and G. Görk (eds.), Pl. Life S.W. and Central Asia 1: 148. = Allium sect. Unicaulea F.O.Khass., 2011. Stapfia 95: 174. (Typus: Allium kotshyi Boiis.)
Typus: Allium lehmannianum Merckl. ex Bunge.

4. Materials and Methods

The object of the study was the endemic species of Kazakhstan A. lehmannianum growing in the northeastern part of Betpakdala, as shown in Figure 1 and Figure 6.

4.1. Distribution Analyses

We compiled distribution maps from the literature and online databases and analyzed herbarium collections (LE, MW, OSBU, AA) and herbarium acronyms according to the Index Herbariorum [27].
Classical botanical (route-reconnaissance, ecological-systematic, ecological-geographical) methods were used in the research. The following fundamental summaries were used to identify the collected material: “Flora of Kazakhstan” [15], “Illustrated Guide to Plants of Kazakhstan” [28], “Guide to Plants of Central Asia and Kazakhstan” [14] and works by authors studying the genus Allium L. [29,30,31,32]. The names of plant species were given by the data from the Plants of the World website [33]. The authorship of species, genera, and families has been critically cross-checked against the information in the International Plant Names Index [34]. The material from the Plantarium website [35] was also used. The QGIS 3.40 program was used for data mapping (https://qgis.org, accessed on 5 January 2025).

4.2. Anatomical Analysis

The specimens for the anatomical studies were collected in vivo during the flowering phase from two populations of the study species located in the western and northern parts of the Betpakdala desert (Central North Turanian subprovince of the North Turanian province of the Iranian-Turanian subregion of the Sahara-Gobi desert region) [36]. Objects for anatomical sections were fixed in 70% alcohol. For the office processing of specimens, 15 × 15 mm histological paraffin was used for fixation. Materials were stained in methylene blue eosin according to May–Grunwald. Cross-sections of samples were made using the automatic microtome (MEDITE M530, MEDITE, Burgdorf, Germany) [37]. Transverse slices were obtained by employing a Levenhuk D740T 5.1 camera using Levenhuk Lite software at ×720, ×1000 magnification. Biometric data were also obtained using this software. The mean number and standard error of biometric data were calculated using Microsoft Excel software using the data analysis function.

4.3. Flow Cytometry

The DNA content was determined via flow cytometry techniques with propidium iodide (PI) staining. Leaves dried with silica gel were used as samples. Samples were chopped using a standard sharp razor blade in LB01 buffer containing PI (50 µg/mL), RNase (50 µg/mL) [38,39], and supplemented with 12 mM sodium thiosulfate and 1% polyvinylpyrrolidone [38]. The nuclear suspension was filtered through a nylon filter with a pore size of 30 μm. Analyses were performed on a Cytoflex cytometer (Beckman Coulter, Inc.) with a 488 nm laser and 610 excitation filter. Analyses were performed on a Cytoflex (Beckman Coulter, Inc., Brea, CA, USA) cytometer. Peaks with at least 1000 nuclei and a CV of less than 5% were analyzed. Histograms were visualized and processed using CytExpert software (Beckman Coulter, Inc., Brea, CA, USA). Descriptive statistics were calculated using XLStat (Addinsoft, Paris, France). An internal standard was used: the Pisum sativum ‘Ctirad’, 2C = 9.09 pg and Vicia faba ‘Inovec’, 2C = 26.9 pg [38,39]. The DNA content was calculated from the ratio of internal standard to sample peaks. The expected ploidy level was interpreted based on published DNA content data in the Kew C-value database for closely related species [40,41]. The angiosperm DNA C-values database (release 9.0, April 2019) (https://cvalues.science.kew.org/) [41] was referred to.

4.4. DNA Extraction, Amplification, and Sequencing

Plant material for DNA extraction was isolated from silica dried leaves collected in the field. Newly sequenced accessions were marked with Am numbers in the trees, and their origin is shown in Table A3. Sequences from the NCBI GenBank are marked with GenBank accession numbers. The total genomic DNA was isolated from silica dried leaves collected in the field using the Diamond Plant DNA Kit (http://diamond-dna.ru/) according to the manufacturer’s instructions and used directly in the PCR amplification. The complete nuclear ribosomal ITS region (ITS1, 5.8S, and ITS2) was amplified using the primers ITS-A [42] and ITS-4 [43]. The chloroplast regions psbA-trnH were amplified using the primers published by Shaw et al. [44] and the trnL intron und trnL - trnF spacer was amplified using the primers published by Taberlet et al. [45]. PCR products were sent for sequencing to Microsynth SeqLab (Göttingen, Germany; www.microsynth.seqlab.de). The sequences from all the individuals were manually edited in Chromas Lite 2.1 (Technelysium Pty Ltd., South Brisbane, Australia) and aligned with ClustalX [46], and the alignment was manually corrected using MEGA 7 [47].

4.5. Phylogenetic Analyses

Sequences from Allium ramosum, A. tuberosum, A. trifurcatum, and A. oreoprasum (A. subgen. Butomissa) and A. vodopjanovae N.Friesen and A. rubens (A. subgen. Rhizirideum) from the third evolution lineage [29,48] were chosen as the outgroup. ITS data sets were analyzed for position identification in the third evolutionary lineage and to find the relationships of A. lehmannianum through parsimony (PAUP) and Bayesian phylogenetic analysis (MrBayes). Fitch parsimony was determined via the heuristic search option in PAUP version 4.0b10 [49] with MULTREES, TBR branch swapping, and 100 replicates of random addition sequences. Gaps were treated as missing data. The consistency index (CI) [50] was calculated to estimate the amount of homoplasy in the character set. The most parsimonious trees returned by the analysis were summarized in one consensus tree using the strict consensus method. Bootstrap analyses using 1000 pseudoreplicates were performed to assess the support (BS) of the clades [51]. Bayesian phylogenetic analyses were also performed using MrBayes 3.1.23 [52]. The sequence evolution model was chosen following the Akaike Information Criterion (AIC) obtained from jModelTest2 [53]. Two independent analyses with four Markov chains were run for 10 million generations, sampling trees every 100 generations. The first 25% of trees were discarded as burn-in. The remaining trees were combined into a single data set, and a majority-rule consensus tree was obtained along with posterior probabilities (PP).

5. Conclusions

The botanical descriptions of both phytocenoses with the participation of Allium lehmannianum showed up to 30 plant species from 16 families. Five species were found in both populations (Artemisia terrae-albae, Astragalus campylotrichus, Bassia prostrata, Oreosalsola arbusculiformis, Tulipa biflora).
The examination of the species via flow cytometry revealed triploidy in the second population, but the DNA amounts of some species of the subgenus Allium have nearly the same amount of DNA. The occurrence of triploid specimens can be explained by possible hybridization.
The analysis of the peculiarities of the anatomy of the vegetative organs of A. lehmannianum showed the derived structures of leaf and stem such as the presence of a thickened outer wall of the epidermis in the leaf; conductive bundles are arranged in a circle in the form of the closed collateral type; and in the anatomical structure of the stem there are internal and external conductive bundles. The main difference between the two populations is the biometric indices of the vegetative organs of A. lehmannianum.
According to the modern phylogenetic tree, Allium lehmannianum belongs to the section Multicaulea, and the section Unicaulea should probably be a synonym of the section Multicaulea. Further studies of the subgenus Allium, which have not yet been studied molecularly, are also needed.

Author Contributions

Conceptualization, D.S.A.; methodology, D.S.A., N.F., B.B.O. and M.V.S.; formal analysis, N.F., P.V.V. and G.M.K.; writing—preparation of the initial draft, D.S.A., P.V.V., G.M.K. and N.F.; editing, N.F., D.S.A., P.V.V. and G.M.K.; author’s supervision, N.F.; project administration, D.S.A. and N.F.; acquisition of funding, P.V.V. All authors have read and agreed to the published version of the manuscript.

Funding

This research has been funded by the Ministry of Science and Higher Education of the Republic of Kazakhstan (Grant No. AP19679078—Studying the species diversity of the ecotone territory of northeastern Betpakdala for preserving the relict gene pool of Kazakhstan arid flora).

Data Availability Statement

All data supporting this study’s findings are available in the main text or Appedixes.

Conflicts of Interest

All authors declare that they have no competing interests and personal relationships and agree on the contents of the paper.

Appendix A

Table A1. List of herbarium specimens viewed in herbarium collections (AA, MW, LE).
Table A1. List of herbarium specimens viewed in herbarium collections (AA, MW, LE).
Herbarium (AA)
Floristic DistrictLabel DataCoordinatesCollectorsDate of Collection
BetpakdalaNorthern shore of Balkhash near Bertys Bay, rocky desertN 46.460001
E 74.044183		Popov M.G.11.VI.1934
Northern Pribalkhashie. Bertys. Rubbly steppe near Novaya Ploschad.N 46.555308
E 74.305633		Zeifert O.A.30.V.1935
Northern Pribalkhashie. Coast of Bertys Bay. Wormwood rubbly desert.N 46.541451
E 74.242282		Rubtsov N.I.30.V.1935
Northern Pribalkhashie. Near the Buru-Baytal settlement. Rubble hills.N 46.667969
E 74.126609		Dmitrieva A.A.26.V.1937
Eastern Betpakdala. Neighbourhood of Myn-Aral settlement. Stony-clay solonchak interfold plain.N 45.443402
E 73.588443		Goloskokov V.P.07.VI.1949
Pribalkhashie, stony slopes of hills along the shore of Balkhash Bay at 225 notch.N 46.537809
E 73.822024		Pavlov N.V.18.V.1951
Jambyl region. 70 km from Priozersk, western part of Betpakdala, northeastern slope.N 46.000268
E 72.792945		Vesselova P.V.,
Kudabayeva G.M.,
Friesen N.V., Osmonali B.B., Urazalina A.S., Abdildanov D.Sh.		28.IV.2024
Karaganda region, northern part of Betpakdala near the Gulshat settlement, stony-rubble slope.N 46.600332
E 74.414978		Vesselova P.V., Kudabayeva G.M., Friesen N.V., Osmonali B.B., Urazalina A.S., Abdildanov D.Sh.29.IV.2024
KaratauZakaratav desert, rubbly areas in the wormwood-bojalych plain between the Uzen and Aksumbe rivers.N 43.630095
E 69.484828		Pavlov N.V.12.V.1939
Jamb.obl., on the slope of red clay hills on the eastern shore of the salt lake Kzyl-Kul.N 43.729897
E 69.432568		Pavlov N.V.02.VI.1948
Shu-IliKhan-Tau steppeN 44.251918
E 73.823705		Baitenov M.S.17.V.1951
Herbarium (MW)
TurgaiSlopes of Khoban-Kulak. Khoban-Kulak, IrsuN 49.285007
E 63.565340		Kryukov I.18.V.1898
Herbarium (LE)
TurgaiTurgai oblast, Kizil-jingilskaya volost, Sarysu river in the lower reaches. Sarysu in the lower reaches. Neighbourhood of the tract. Tes-Bulak thickets of biurguns and NanaphytonN 49.175696
E 63.519753		Krasneshnikov I.M.03.VI. 1914
KaratauNorthern slope of Northern Karatau slope of Balasauskandyk mountainsN 43.364328
E 69.787435		Kamelin R.V., Mikhailova M.A., Mishenkova A.P., Safronova I.N., Soloviev V.I.14.V.1977

Appendix B

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Figure A1. A. lehmannianum (Herbarium AA).
Figure A1. A. lehmannianum (Herbarium AA).
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Appendix C

Table A2. Species community composition with the participation of the studied species.
Table A2. Species community composition with the participation of the studied species.
№SpeciesPopul.Life FormPhenoph.EcotypeNotes
I II
I. Amaryllidaceae J.St.-Hil.
1Allium lehmannianum++PerennialBl. V–VIXerophyteEndeme
II. Amaranthaceae Juss.
2Anabasis truncata (Schrenk) Bunge+ PerennialBl. VIIXerophyte
3Anabasis salsa (C.A. Mey.) Benth. ex Volkens+ Semi-shrubBl. VIIXerophyte
4Bassia prostrata (L.) Beck++Half-shrubBl. and fr.
VII–IX		Xerophyte
5Ceratocarpus arenarius L. +AnnualBl. and fr.
V–VII		Xerophyte
6Nanophyton erinaceum (Pall.) Bunge.+ ShrubBl. VIIXerophyte
7Oreosalsola arbusculiformis (Drobow) Sennikov (Salsola arbusculiformis Drobow.)++ShrubBl.V–VIIIXerophyte
8Krascheninnikovia ceratoides (L.) Gueldenst., +ShrubBl. VIII–IXXerophyte
III. Asphodelaceae Juss.
9Eremurus cristatus+ PerennialBl. V–VIXero-mesophyte
IV. Asteraceae Bercht. & J.Presl
10Artemisia terrae-albae++Half-shrubBl. VIII–XMesophyte
V. Boraginaceae Juss.
11Rochelia retorta (Pall. Lipsky)+ AnnualBl. IV–VXerophyte
12Lappula spinocarpos (Forssk.) Asch. +AnnualBl. IV–VXerophyte
VI. Brassicaceae Burnett
13Descurainia sophia (L.) Webb ex Prantl +AnnualBl. V–VIIIXero-mesophyte
14Megacarpaea megalocarpa (Fisch. ex DC.) B. Fedtsch. +PerennialBl. IV–VXerophyteRelict
15Meniocus linifolius (Stephan ex Willd.) DC. +AnnualBl. III–IVXero-mesophyte
VII. Cyperaceae Juss.
16Carex pachystylis J. Gay, +PerennialBl. IV–VIMesophyte
17Carex physodes M. Bieb. +PerennialBl. IV–VIMesophyte
VIII. Ephedraceae Dumort.
18Ephedra distachya L.+ ShrubBl. V–VI
Fr. VII–VIII		Xerophyte
IX. Euphorbiaceae Juss.
19Euphorbia rapulum Kar. and Kir.+ PerennialBl. IV–VXerophyte
X. Fabaceae Lindl.
20Astragalus campylotrichus Bunge++AnnualBl. IV–V
Fr. V–VI		Xerophyte
XI. Iridaceae Juss.
21Iris tenuifolia Pall. +PerennialBl. IV–VXerophyte
XII. Liliaceae Juss.
22Fritillaria karelinii (Fisch. ex D.Don) Baker +PerennialBl. III–V
Fr. V–VI		Xero-mesophyte
23Tulipa biflora Pall. (Tulipa buhseana Boiss.)++PerennialBl. IV–VXero-mesophyte
24Tulipa alberti Regel+ PerennialBl. IV–VXero-mesophyterare species
XIII. Poaceae Barnhart
25Festuca valesiaca Gaudin+ PerennialBl. V–VIXero-mesophyte
26Eremopyrum bonaepartis (Spreng.) Nevski+ AnnualBl. IV–VXero-mesophyte
27Stipa caucasica Schmalh. +PerennialBl.IV–VIXerophyte
XIV. Polygonaceae Juss.
28Rheum tataricum L. f+ PerennialBl. IV–V
Fr. V–VI		Xero-mesophyte
XV. Ranunculaceae Juss.
29Ranunculus testiculatus Crantz +AnnualBl. III–VMesophyte
XVI. Zygophyllaceae R.Br.
30Zygophyllum pinnatum Cham.+ PerennialBl. IV–V
Fr. V–VIII		Xerophyte
The + sign indicates the presence of a species in a population.
Table A3. Vouchers of the species studied.
Table A3. Vouchers of the species studied.
AccessionSpeciesOriginLat_LonITSpsbA-trnHtrnL-trnF
Am1501A. lehmannianumKZ: West Betpakdala46.000268 N, 72.792945 EPQ890140PQ894028PQ894030
Am1501_2A. lehmannianumKZ: West Betpakdala46.000268 N, 72.792945 EPQ890141
Am1514_1A. lehmannianumKZ: East Betpakdala46.600332 N, 74.414978 EPQ890142PQ894029PQ894031
Am1514_1A. lehmannianumKZ: East Betpakdala46.600332 N, 74.414978 EPQ890143
Am1514_3A. lehmannianumKZ: East Betpakdala46.600332 N, 74.414978 EPQ890144
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Figure A2. Distribution of Allium kotschyi (yellow dots) (GBIF https://www.gbif.org/species/2856108. accessed on 5 January 2025). (Type: Iran, Kuh-e Dinar Mount, 30.952842195, 51.4375855).
Figure A2. Distribution of Allium kotschyi (yellow dots) (GBIF https://www.gbif.org/species/2856108. accessed on 5 January 2025). (Type: Iran, Kuh-e Dinar Mount, 30.952842195, 51.4375855).
Plants 14 01113 g0a2

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Plants 14 01113 g001
Figure 1. Distribution of A. lehmannianum according to our collections and herbarium data.
Figure 1. Distribution of A. lehmannianum according to our collections and herbarium data.
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Plants 14 01113 g002
Figure 2. Family spectrum of species composition of two communities with participation of A. lehmannianum.
Figure 2. Family spectrum of species composition of two communities with participation of A. lehmannianum.
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Plants 14 01113 g003
Figure 3. Comparative anatomical structure of the studied species: (A) transverse section of the leaf plate (×720), (B) structure of the conductive bundle of the leaf plate (×1000), (C) cross-section of the stem (×720), (D) structure of the conductive bundle of the stem (×1000). K—cuticle, E—epidermis, PM—polysaccharide mesophyll, SM—spongy mesophyll, SC—sclerenchyma, Xy—xylem (outer), Ph—phloem (outer), PX—main parenchyma.
Figure 3. Comparative anatomical structure of the studied species: (A) transverse section of the leaf plate (×720), (B) structure of the conductive bundle of the leaf plate (×1000), (C) cross-section of the stem (×720), (D) structure of the conductive bundle of the stem (×1000). K—cuticle, E—epidermis, PM—polysaccharide mesophyll, SM—spongy mesophyll, SC—sclerenchyma, Xy—xylem (outer), Ph—phloem (outer), PX—main parenchyma.
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Plants 14 01113 g004
Figure 4. Examples of flow cytometric histograms of the A. lehmannianum samples (log scale). (A) diploid cytotype with P. sativum as internal standard; (B) diploid cytotype with V. faba as internal standard; (C) triploid cytotype with V. faba as internal standard.
Figure 4. Examples of flow cytometric histograms of the A. lehmannianum samples (log scale). (A) diploid cytotype with P. sativum as internal standard; (B) diploid cytotype with V. faba as internal standard; (C) triploid cytotype with V. faba as internal standard.
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Plants 14 01113 g005
Figure 5. Phylogenetic nrITS tree of the subgenus Allium of genus Allium. Numbers by nodes represent bootstrap support (1000 replicates) and Bayesian probabilities. The joint presence of Bayesian probabilities over 0.98 and bootstrap support over 95% is indicated with a black dot. For the origin of samples without GenBank accession numbers, see Table A2.
Figure 5. Phylogenetic nrITS tree of the subgenus Allium of genus Allium. Numbers by nodes represent bootstrap support (1000 replicates) and Bayesian probabilities. The joint presence of Bayesian probabilities over 0.98 and bootstrap support over 95% is indicated with a black dot. For the origin of samples without GenBank accession numbers, see Table A2.
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Figure 6. Object of study A. lehmannianum.
Figure 6. Object of study A. lehmannianum.
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Table 1. Biometric data of anatomical sections of leaf blade of A. lehmannianum (μm).
Table 1. Biometric data of anatomical sections of leaf blade of A. lehmannianum (μm).
KEPMXyPhPX
Population 1
21 ± 3.3621.7 ± 3.3979.85 ± 9.17 × 16.21 ± 2.30 8.80 ± 1.468.65 ± 2.2542.36 ± 4.13
Population 2
21 ± 3.0223.30 ± 3.0590.23 ± 9.51 × 20 ± 2.810.27 ± 2.275.82 ± 0.9558.74 ± 4.44
Table 2. Biometric data of anatomical sections of the stem of A. lehmannianum (μm).
Table 2. Biometric data of anatomical sections of the stem of A. lehmannianum (μm).
KESMSCXyPhXy2Ph2PX
Population 1
18 ± 3.3624.19 ± 3.4523.35 ± 3.0924.4 ± 2.048.97 ± 1.554.76 ± 1.4612.89 ± 4.159.04 ± 3.3651.18 ± 4.45
Population 2
21 ± 3.0219.55 ± 3.1621.21 ± 4.1919.59 ± 5.349.22 ± 1.557.59 ± 1.8918.22 ± 3.4412.74 ± 2.1451.27 ± 4.43
Legend: K—cuticle, E—epidermis, SM—spongy mesophyll, SC—sclerenchyma, Xy—xylem (outer), Ph—phloem (outer), Xy2—xylem (inner), Ph2—phloem (inner), PX—main parenchyma.
Table 3. DNA content of the studied A. lehmannianum samples.
Table 3. DNA content of the studied A. lehmannianum samples.
SpeciesLocalityMean 2C, pgSD, pgCV1C, GbpExpected Ploidy
A. lehmannianum (2×)Pop 135.4680.4051.14%34.6882
A. lehmannianum (2×)Pop 235.1040.2920.83%34.3322
Mean 2C for diploid cytotype35.3230.3931.11%
A. lehmannianum (3×)Pop 250.2120.4170.83%49.1073
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Abdildanov, D.S.; Vesselova, P.V.; Kudabayeva, G.M.; Osmonali, B.B.; Skaptsov, M.V.; Friesen, N. Endemic of Kazakhstan Allium lehmannianum Merckl. Ex Bunge and Its Position Within the Genus Allium. Plants 2025, 14, 1113. https://doi.org/10.3390/plants14071113

AMA Style

Abdildanov DS, Vesselova PV, Kudabayeva GM, Osmonali BB, Skaptsov MV, Friesen N. Endemic of Kazakhstan Allium lehmannianum Merckl. Ex Bunge and Its Position Within the Genus Allium. Plants. 2025; 14(7):1113. https://doi.org/10.3390/plants14071113

Chicago/Turabian Style

Abdildanov, Daulet Sh., Polina V. Vesselova, Gulmira M. Kudabayeva, Bektemir B. Osmonali, Michail V. Skaptsov, and Nikolai Friesen. 2025. "Endemic of Kazakhstan Allium lehmannianum Merckl. Ex Bunge and Its Position Within the Genus Allium" Plants 14, no. 7: 1113. https://doi.org/10.3390/plants14071113

APA Style

Abdildanov, D. S., Vesselova, P. V., Kudabayeva, G. M., Osmonali, B. B., Skaptsov, M. V., & Friesen, N. (2025). Endemic of Kazakhstan Allium lehmannianum Merckl. Ex Bunge and Its Position Within the Genus Allium. Plants, 14(7), 1113. https://doi.org/10.3390/plants14071113

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