Morpho-anatomical features and adaptive traits of Eremurus (Asphodelaceae) species in the Almaty Region, KazakhstanAbstract
A comprehensive study of the morphological, anatomical, and ecological characteristics of six Eremurus M. Bieb. species (E. robustus, E. tianschanicus, E. fuscus, E. cristatus, E. inderiensis, and E. anisopterus) was conducted in the Almaty Region and the Northern Tien Shan. The study aimed to identify interspecific differences and adaptive traits formed under arid and mountainous environmental conditions. Field material was collected during expeditions in 2025. For each species, 10 individuals from natural populations were examined, and at least 30 quantitative measurements were performed for each morphological and anatomical trait. Morphological analysis revealed considerable variability in plant habitus, leaf morphology, inflorescence architecture, and the dimensions of the perianth and fruits. Fruit diameter (M ± SD) was analyzed as a reproductive morphological trait with potential taxonomic relevance.One-way analysis of variance (ANOVA) demonstrated a statistically significant effect of the factor “species” on plant height, leaf width, perianth length, and fruit diameter, confirming their diagnostic and adaptive significance. All studied species exhibit pronounced xeromorphic features, including a single-layered epidermis with a well-developed cuticle, sunken stomata, differentiated mesophyll, and water-storage cells. Quantitative anatomical parameters vary considerably among species. The most pronounced xeromorphic and water-accumulating traits were recorded in E. anisopterus and E. inderiensis, which inhabit sandy and highly arid environments. The novelty of the study lies in the first integrated comparative morpho-anatomical assessment of six co-occurring Eremurus species in southeastern Kazakhstan, demonstrating the ecological and taxonomic significance of anatomical traits in arid mountain ecosystems. The practical significance of the study lies in the identification of diagnostic morpho-anatomical traits that can be used for species identification, ecological assessment, biodiversity monitoring, and conservation planning of Eremurus species in arid and mountain ecosystems of southeastern Kazakhstan.
Keywords:
Eremurus; anatomy; morphology; palisade mesophyll; water-storage cells; Northern Tien Shan
Resumo
Foi realizado um estudo abrangente das características morfológicas, anatômicas e ecológicas de seis espécies de Eremurus M. Bieb. (E. robustus, E. tianschanicus, E. fuscus, E. cristatus, E. inderiensis e E. anisopterus) na região de Almaty e no Norte do Tien Shan. O objetivo do estudo foi identificar diferenças interespecíficas e traços adaptativos formados sob condições ambientais áridas e montanhosas. O material de campo foi coletado durante expedições em 2025. Para cada espécie, 10 indivíduos de populações naturais foram examinados, e pelo menos 30 medições quantitativas foram realizadas para cada característica morfológica e anatômica. A análise morfológica revelou considerável variabilidade no hábito da planta, na morfologia das folhas, na arquitetura da inflorescência e nas dimensões do perianto e dos frutos. O diâmetro do fruto (M ± DP) foi analisado como uma característica morfológica reprodutiva com potencial relevância taxonômica. A análise de variância de um fator (ANOVA) demonstrou um efeito estatisticamente significativo do fator “espécie” sobre a altura da planta, a largura da folha, o comprimento do perianto e o diâmetro do fruto, confirmando sua importância diagnóstica e adaptativa. Todas as espécies estudadas apresentam características xeromórficas pronunciadas, incluindo epiderme unisseriada com cutícula bem desenvolvida, estômatos afundados, mesófilo diferenciado e células de armazenamento de água. Os parâmetros anatômicos quantitativos variam consideravelmente entre as espécies. As características xeromórficas e de acúmulo de água mais pronunciadas foram registradas em E. anisopterus e E. inderiensis, que habitam ambientes arenosos e altamente áridos. A novidade do estudo reside na primeira avaliação comparativa integrada morfoanatômica de seis espécies coexistentes de Eremurus no sudeste do Cazaquistão, demonstrando a importância ecológica e taxonômica dos caracteres anatômicos em ecossistemas montanhosos áridos. O significado prático do estudo reside na identificação de características morfoanatômicas diagnósticas que podem ser usadas para identificação de espécies, avaliação ecológica, monitoramento da biodiversidade e planejamento da conservação de espécies de Eremurus em ecossistemas áridos e montanhosos do sudeste do Cazaquistão.
Palavras-chave:
Eremurus; anatomia; morfologia; mesófilo paliçádico; células de armazenamento de água; Norte do Tien Shan
1. Introduction
The genus Eremurus M. Bieb. (Asphodelaceae) includes approximately 70 species of perennial herbaceous plants distributed mainly in the mountain-steppe and foothill regions of Central and East Asia, with several representatives extending to Crimea, the Caucasus, and Western Siberia. Species of the genus are well known for their ecological plasticity and adaptation to arid and semi-arid environments. Many species inhabit dry slopes, foothill deserts, and mountain-steppe ecosystems, where plants are exposed to high solar radiation, seasonal drought, and strong temperature fluctuations. Due to their large inflorescences and ornamental value, Eremurus species have also been introduced into botanical gardens and horticultural collections in Europe and Asia since the nineteenth century (Baitenov, 2001; Naderi et al., 2014; Bekkulova and Mukumov, 2021; Makhmudjanov et al., 2019; Jang et al., 2024).
Species of Eremurus possess a number of distinctive morphological traits that facilitate their identification. These include a leafless flowering scape bearing a large racemose inflorescence often composed of numerous flowers, as well as a well-developed rhizomatous root system adapted to seasonal drought conditions. On the basis of floral morphology, the genus is traditionally divided into two subgenera: subgenus Eremurus, characterized by tubular or campanulate flowers of light brown or greenish coloration, and subgenus Henningia, which includes species with radially symmetrical white, pink, or yellow flowers (Makhmudjanov et al., 2023, 2025). This subdivision reflects morphological and evolutionary differentiation within the genus.
Previous studies on Eremurus have addressed various aspects of its biology, including taxonomy, morphology, reproductive biology, and ornamental potential. Morphological diversity and reproductive characteristics of several species have been investigated in different regions of Eurasia, demonstrating significant variation in plant architecture, seed productivity, and floral biology. For example, studies conducted in Crimea revealed differences in flowering phenology and seed formation among several species, whereas research in China showed functional differentiation of flowers within the inflorescence in E. anisopterus, which contributes to reproductive efficiency. Investigations in Iran and Central Asia have also demonstrated considerable morphological variation associated with geographic and environmental factors (Lysyakova et al., 2009; Schiappacasse et al., 2013; Kumari and Saggoo, 2016; Hadizadeh et al., 2021; Jannathan et al., 2014; Makhmudjanov et al., 2022; Sayed et al., 2023).
In addition to morphological variability, recent research has emphasized the importance of anatomical and physiological traits in understanding plant adaptation to arid environments. Anatomical studies of several Eremurus species have revealed the presence of xeromorphic features such as a single-layered epidermis with a developed cuticle, sunken stomata, and mesophyll composed of large parenchymatous cells that may function in water storage. Such structural characteristics are considered important adaptive mechanisms allowing plants to tolerate water deficit and high irradiance. For example, studies of E. himalaicus, E. robustus, and E. stenophyllus have demonstrated species-specific differences in leaf anatomy and stomatal structure associated with ecological conditions (Mushtaq et al., 2016; Bahrim et al., 2017, 2020; Duschanova et al., 2023; Mokhtar et al., 2024).
Despite the growing interest in the genus, comparative morpho-anatomical studies of Eremurus species in southeastern Kazakhstan remain limited. The flora of the Almaty Region and the Northern Tien Shan includes several species of the genus that occur in contrasting ecological conditions, ranging from foothill steppe habitats to sandy and rocky mountain slopes. However, the structural adaptations that allow these species to persist under different environmental conditions remain insufficiently studied, particularly with respect to quantitative anatomical traits.
Quantitative anatomical metrics provide valuable information for assessing interspecific differentiation and ecological adaptation. Measurements of epidermal thickness, mesophyll structure, vascular bundle dimensions, and water-storage tissues allow the identification of structural features associated with xeromorphic adaptation and can complement traditional morphological characters used in taxonomy.
Therefore, the present study aims to conduct a comparative analysis of morphological and anatomical traits of six Eremurus species occurring in the Almaty Region and the Northern Tien Shan. The objective of the research is to identify common and species-specific structural features, evaluate their adaptive significance under arid and mountainous environmental conditions, and assess the taxonomic and ecological value of anatomical characteristics within the genus.
2. Materials and Methods
2.1. Study objects
The objects of the study were representatives of the genus Eremurus M. Bieb. (Asphodelaceae) occurring within the Almaty Region and differing in ecological preferences and levels of morpho-anatomical organization. The study included six species: Eremurus robustus Regel, E. tianschanicus Pazij & Vved. ex Pavlov, E. fuscus (O. Fedtsch.) Regel, E. cristatus Vved., E. inderiensis (Boiss.) Regel, and E. anisopterus (Kar. & Kir.) Regel (Figure 1).
Research objects: (A) Eremurus cristatus; (B) E. inderiensis; (C) E. fuscus; (D) E. anisopterus; (E) E. tianschanicus; (F) E. robustus.
The selection of these species was based on their wide representation in the flora of the Almaty Region, differences in ecological niches (mountain, foothill, and arid habitats), and pronounced variability in morphological and anatomical traits. These characteristics make them suitable model taxa for identifying adaptive strategies of the genus Eremurus under regional environmental conditions.
2.2. Study area
The study area covered natural landscapes of the Almaty Region in southeastern Kazakhstan, including foothill and mountain systems of the Northern Tien Shan (Zailiysky Alatau and Kungey Alatau ranges) as well as adjacent plains and semi-desert territories (Figure 2). The elevation of the collection sites ranged approximately from 500 to 2200 m above sea level, encompassing foothill steppe zones, rocky slopes, mountain meadows, and arid habitats.
Map of the study area showing the location of the Almaty Region within southeastern Kazakhstan.
The regional climate is strongly continental, characterized by pronounced seasonal temperature fluctuations and uneven precipitation distribution. The mean annual temperature ranges from approximately 7-9 °C in foothill areas to 3-5 °C in mountain zones, while annual precipitation varies from about 250-350 mm in lowland and foothill territories to 500-700 mm in mountainous areas. Such environmental heterogeneity creates diverse ecological conditions that influence plant morphology and anatomical structure.
The studied species occupy different habitat types within the region. Eremurus robustus and E. tianschanicus mainly occur on mountain slopes and foothill meadows with relatively higher soil moisture. E. fuscus and E. cristatus are typically found on steppe slopes and rocky substrates, while E. inderiensis and E. anisopterus are associated with more arid habitats, including sandy and gravelly soils of foothill and semi-desert landscapes.
The combination of altitudinal gradients, diverse substrates, and contrasting moisture regimes makes the Almaty Region a representative model area for studying morpho-anatomical variability and ecological adaptation in species of the genus Eremurus.
2.3. Field sampling design
Field sampling was carried out during botanical expeditions in 2025 across different parts of the Almaty Region and the Northern Tien Shan. The sampling design aimed to represent the ecological diversity of habitats in which species of the genus Eremurus occur.
The unit of biological replication was the individual plant (n = 10 per species), while repeated measurements (≥30) were treated as technical replicates used to estimate within-individual variability. Statistical analyses were performed using mean values per individual to ensure independence.
For each studied species, one natural population was selected within its typical habitat, and 10 generative individuals were sampled from each population. Individuals were selected using a randomized approach within the population area, maintaining a minimum distance of 3-5 m between plants to avoid sampling of clonally related individuals. This approach allowed us to obtain representative material reflecting the morphological and anatomical variability of each species under natural conditions (Kubentayev et al., 2022; Kenesbay et al., 2025).
The sampling design allows comparison of morpho-anatomical traits among species under specific local environmental conditions; however, it does not capture the full range of intraspecific variability across the species’ distribution.
During fieldwork, geographic coordinates (WGS84), elevation above sea level, habitat characteristics, and associated vegetation were recorded for each sampling site. The studied populations were located in different ecological conditions, including foothill steppe slopes, mountain meadows, rocky substrates, and sandy arid habitats (Danilova et al., 2025; Islamgulova et al., 2025; Orazov et al., 2025; Osmonali et al., 2025).
Collected plant material was used for both morphological measurements and anatomical analyses. From each individual, vegetative organs (leaves and stems) and generative structures (flowers and fruits) were sampled. For morphometric and anatomical analyses, at least 30 quantitative measurements were performed for each trait, ensuring sufficient replication for statistical analysis.
This sampling strategy allowed us to compare morpho-anatomical characteristics among species and to assess their adaptive structural traits under contrasting environmental conditions.
2.4. Anatomical methods
In addition to morphological traits, selected anatomical parameters (epidermal thickness, palisade mesophyll thickness, and water-storage cell diameter) were analyzed using one-way ANOVA under the same statistical framework.
For anatomical investigations, plant samples were prepared following standard histological techniques. Fresh plant material was fixed in a 70% ethanol solution, which ensured the preservation of tissue structures and prevented cellular degradation during further processing.
Transverse sections were prepared from the middle part of fully developed leaves and from stem internodes of generative shoots. For each individual plant, 3-5 anatomical sections were obtained for both organs to ensure representative observation of tissue structures. The samples were preliminarily frozen and subsequently embedded in paraffin, which was poured into special molds measuring 15 × 15 mm.
The solidified paraffin blocks were sectioned using a semi-automatic rotary microtome (MEDITE M530, Burgdorf, Germany). Transverse sections 40 μm thick were obtained and mounted on microscope slides. To enhance tissue differentiation, sections were stained with safranin solution, which allowed clearer visualization of cell walls and vascular tissues.
Microscopic examination was performed using a digital microscope (Levenhuk Zoom&Joy, Hong Kong, China) equipped with a Levenhuk D740T 5.1 camera, enabling the acquisition of high-resolution images. Image acquisition, processing, and morphometric measurements were conducted using LevenhukLite software, version 4.12.28273 (Turgara et al., 2026).
Prior to morphometric analysis, the measurement system was calibrated using a stage micrometer, which allowed all anatomical parameters to be expressed in micrometers (µm). For each anatomical trait, at least 30 quantitative measurements were performed across different sections to ensure statistical reliability.
To minimize measurement error, sections with mechanical damage or deformation were excluded from analysis. Only well-preserved and clearly visible tissues were used for morphometric measurements. The obtained biometric data were subjected to statistical analysis, including the calculation of mean values, standard deviations, and other statistical parameters using the Data Analysis module of Microsoft Excel (Islamgulova et al., 2025). This methodological approach ensured the reliability and comparability of anatomical measurements among the studied Eremurus species.
2.5. Statistical analysis
Statistical analysis was performed to identify interspecific differences in morphological traits among representatives of the genus Eremurus. The factor variable was species (Eremurus robustus, E. tianschanicus, E. fuscus, E. cristatus, E. inderiensis, E. anisopterus), while the dependent variables included quantitative morphological traits: plant height, leaf width, perianth length, and fruit diameter.
For each species, 10 individuals were analyzed (n = 10). Prior to statistical analysis, the dataset was tested for compliance with the assumptions of parametric statistics. Normality of distribution was assessed using the Shapiro-Wilk test, and homogeneity of variances was evaluated using Levene’s test (Sumbembayev et al., 2023, 2025).
If these assumptions were satisfied, one-way analysis of variance (ANOVA) was applied to determine statistically significant differences among species. When significant differences were detected (p < 0.05), Tukey’s HSD post-hoc test was used to identify pairwise differences between species.
All statistical analyses were performed using Statistica software, version 13.5 (TIBCO Software Inc., USA). For each analyzed trait, the ANOVA results are presented as F-values, degrees of freedom (df), and significance levels (p). In addition, effect size (η2) was calculated to estimate the contribution of the species factor to the observed variation in morphological traits (Kubentayev et al., 2022; Sumbembayev et al., 2023, 2025).
3. Results
As a result of expeditionary surveys conducted in 2025 with the aim of collecting plant material for morphological and anatomical analyses, specimens of six species of the genus Eremurus were collected from various natural and administrative districts of the Almaty Region. The field studies covered mountainous, foothill, and arid landscapes of the region, which made it possible to document the spatial distribution of the species and to ensure the representativeness of the studied material. For each species, the geographical coordinates of the collection sites were recorded and habitat conditions were updated, providing a basis for subsequent analysis of morpho-anatomical characteristics and adaptive traits (Table 1; Figure 3).
3.1. Morphological characteristics of the studied species
Representatives of the genus Eremurus in the flora of the Almaty Region demonstrate considerable morphological diversity associated with their ecological distribution and adaptive strategies. The studied species differ in plant height, leaf morphology, inflorescence structure, and dimensions of generative organs (Table 2).
Eremurus robustus Regel is the largest species among those studied, reaching 100-200 cm in height. It is characterized by a robust glabrous stem, wide strap-shaped leaves (3-8 cm), and a large racemose inflorescence. The perianth is bell-shaped, 15-17 mm long, pink or occasionally white (Figure 4).
Morphological features of Eremurus robustus: (A) general view of the plant; (B) fruit on the pedicel; (C, E) fruits; (D) flowers in the inflorescence; (F, J) flower; (H) flowers (individual elements of the perianth); (G) inflorescence in the flowering phase; (I) part of the inflorescence with flowers.
Eremurus tianschanicus reaches 50-150 cm in height and differs from the previous species by its narrower leaves (0.5-1.2 cm) and a dense cylindrical inflorescence. The perianth is pale pink with a yellow base and measures 10-12 mm in length (Figure 5).
Morphological features of Eremurus tianschanicus: (A) general view of the plant; (B) anther of the stamens; (C) part of the inflorescence with flowers; (D) fruit on the pedicel; (E, G) inflorescence in the flowering phase; (F) anther; (H, J) fruits; (I) part of the inflorescence with open flowers.
Eremurus fuscus is a medium-sized species (70-100 cm tall) with leaves 1.5-4 cm wide and a dense cylindrical inflorescence. The perianth is narrowly bell-shaped, 10-12 mm long, light yellow or fawn, gradually turning brown after flowering (Figure 6).
Morphological features of Eremurus fuscus: (A) individual flower with well-defined stamens; (B) inflorescence in the flowering phase; (C) flowering inflorescence (general view); (D) flowers in the inflorescence (lateral view); (E) inflorescence; (F) individual flower (detail of the perianth and stamens); (G) plants in their natural habitat (general view); (H) stamens with anthers; (I) inflorescence with fruits; (J) stamen (detail of the anther); (K) inflorescence after flowering.
Eremurus cristatus is a relatively smaller species (50-70 cm tall) with linear glaucous leaves 1-2 cm wide and a compact raceme. The perianth is narrowly bell-shaped, 8-10 mm long, with a characteristic brown stripe on the outer surface (Figure 7).
Morphological features of Eremurus cristatus: (A) inflorescence with fruits (general view); (B) individual flower in the inflorescence; (C, D) inflorescence; (E) fruit; (F) flower with stamens and pistil; (G) flowering inflorescence; (H) inflorescence with fruits; (I) dried inflorescence with fruits; (J) individual flower.
Eremurus inderiensis reaches 80-130 cm in height and is characterized by linear to linear-lanceolate leaves 0.6-2 cm wide and a dense cylindrical inflorescence. The perianth is narrowly bell-shaped, approximately 10 mm long, with a purple coloration and a green median stripe (Figure 8).
Morphological features of Eremurus inderiensis: (A) inflorescence with fruits (general view); (B) inflorescence in the flowering phase; (C) fruits; (D) inflorescence after flowering; (E) general view; (F) flowers in the inflorescence (lateral view); (G) flower; (H) plants in their natural habitat; (I) inflorescence (general view); (J) flowering inflorescence (enlarged fragment).
Eremurus anisopterus is the most compact species (30-70 cm tall), with very narrow basal leaves (0.4-0.6 cm wide) and a relatively short, spreading inflorescence. The perianth is bell-shaped, pale pink or white, reaching up to 20 mm in length (Figure 9).
Morphological features of Eremurus anisopterus: (A) flower; (B) flowers in the inflorescence; (C) individual flower; (D) inflorescence in the flowering phase (upper part); (E) flowering plant in its natural habitat (general view); (F) fruits with preserved perianth; (G) inflorescence with fruits; (H) individual flower; (I) plant in the flowering phase (general view); (J) inflorescence with fruits.
The main morphological differences among the studied species are reflected in plant height, leaf width, and the size of generative organs. Quantitative comparison of these traits, including fruit diameter (M ± SD), is presented in Table 2, and the statistical significance of interspecific differences was evaluated using one-way ANOVA with Tukey’s HSD test.
The quantitative trait presented as fruit diameter was measured as the maximum transverse diameter of mature capsules in all studied species. This parameter was selected as an integral reproductive character for comparative analysis. Measurements were performed on mature, fully developed fruits collected from 10 individuals per species, with at least 30 measurements analyzed for each taxon. Considerable interspecific variation was observed in capsule size, with the smallest mean values recorded in E. fuscus and E. cristatus, intermediate values in E. robustus and E. tianschanicus, and the largest capsules in E. inderiensis and especially E. anisopterus. Because the table summarizes capsule dimensions rather than seed size, these values are biologically comparable within the genus and reflect real differences in reproductive morphology among the studied species.
The markedly larger capsule diameters in E. inderiensis and E. anisopterus reflect true interspecific differences in fruit morphology; however, these values represent capsule size rather than seed dimensions and therefore should not be directly compared with seed-based metrics reported in other studies.
The high variability observed in water-storage cell diameter, particularly in E. robustus, likely reflects a combination of biological heterogeneity among cells and variation across tissue sections rather than measurement error.
3.2. Leaf anatomical structure
The leaf anatomy of the studied Eremurus species is characterized by pronounced xeromorphic features. The leaf blade is covered with a single-layered epidermis with a well-developed cuticle, while the stomata are mostly sunken below the epidermal level, which reduces water loss and enhances resistance to arid environmental conditions.
The mesophyll is differentiated into palisade and spongy parenchyma. The palisade layer consists of elongated, tightly arranged cells that provide efficient photosynthetic activity. The central part of the mesophyll is composed of large parenchymatous cells that function as water-storage cells, allowing the accumulation and retention of moisture under conditions of limited water availability.
The conducting system is represented by well-developed vascular bundles distributed throughout the leaf tissue, ensuring effective transport of water and assimilates. The presence of differentiated mesophyll, sunken stomata, and water-storage cells reflects structural adaptations of Eremurus species to xeric habitats (Figure 10).
Anatomical structure of leaf cross-sections in six species of the genus Eremurus: (A, B) Eremurus robustus; (C, D) E. tianschanicus; (E, F) E. cristatus; (G, H) E. fuscus; (I, J) E. anisopterus; (K, L) E. inderiensis. Abbreviations: c – cuticle; eh – epidermal hair; ep – epidermis; sa – stomatal apparatus; pal – palisade mesophyll; vb – vascular bundle; wc – water-storage cells. Images represent typical transverse sections obtained from multiple replicates (10 individuals per species).
Morphometric measurements of the main anatomical tissues were performed for six species of the genus Eremurus in order to identify interspecific differences in the structure of leaves and stems (Figure 11). The analyzed parameters included epidermal thickness, palisade mesophyll thickness, and diameter of water-storage cells in the leaf (Table 3), as well as epidermal thickness, diameter of primary cortical cells, and diameter of water-storage cells in the stem (Table 4; Figure 11).
Anatomical structure of stem cross-sections in six species of the genus Eremurus: (A, B) Eremurus robustus; (C, D) E. tianschanicus; (E, F) E. cristatus; (G, H) E. fuscus; (I, J) E. anisopterus; (K, L) E. inderiensis. Abbreviations: c – cuticle; ep – epidermis; en – endodermis; cx – primary cortex; par – parenchyma; vb – vascular bundle; xy – xylem; ph – phloem; wc – water-storage cells; eh – epidermal hair. Images represent typical transverse sections obtained from multiple replicates (10 individuals per species).
Anatomical characteristics of leaf tissues in species of the genus Eremurus (µm; n = 10 individuals, ≥30 measurements per trait).
Anatomical characteristics of stem tissues in species of the genus Eremurus (µm; n = 10 individuals, ≥30 measurements per trait).
All measurements were performed in micrometers (µm). For each species, 10 individuals were analyzed, and at least 30 measurements were taken for each anatomical parameter. The data are presented as mean ± standard deviation (M ± SD).
3.3. Ecological characteristics of the studied species
Field observations (present study) indicate that the studied Eremurus species occupy habitats that differ markedly in substrate type, moisture availability, and elevation. The investigated populations were recorded in foothill steppes, rocky slopes, mountain meadows, and sandy habitats, reflecting a wide ecological amplitude of the genus within southeastern Kazakhstan.
Eremurus robustus and E. tianschanicus were mainly observed in foothill and lower mountain habitats, including meadow slopes and steppe communities with relatively deeper soils and moderate moisture availability. In contrast, E. cristatus and E. fuscus occurred predominantly on rocky or gravelly slopes with well-drained substrates and high solar exposure. The species E. anisopterus and E. inderiensis were associated with more arid habitats, including sandy and semi-desert landscapes characterized by loose substrates and limited water availability.
These habitat differences correspond to variations in morphological and anatomical traits observed among species, particularly in leaf structure and the development of water-storage tissues.
According to published literature, species of the genus Eremurus are generally adapted to xerophytic environments of Central Asia, where they occur in open habitats with high insolation and seasonal water deficit. Many species are confined to mountain-steppe, foothill, and desert landscapes, demonstrating a high degree of ecological specialization.
Literature sources indicate that E. cristatus is typically associated with loess steppes and rocky foothill slopes, whereas E. fuscus occurs on mountain slopes with steppe vegetation and gravelly substrates. E. tianschanicus is mainly confined to loess foothills and dry mountain slopes, while E. robustus shows a broader ecological amplitude and may occur in foothill meadows and mountain slopes with relatively higher moisture. The desert species E. inderiensis and E. anisopterus are characteristic of sandy desert ecosystems, where they form part of psammophytic plant communities.
The combination of field observations and literature data confirms that species of the genus Eremurus occupy a gradient of ecological conditions, ranging from relatively humid foothill habitats to highly arid sandy landscapes. Such ecological differentiation is consistent with the morphological and anatomical adaptations identified in the present study.
4. Discussion
4.1. Morphological features
The morphological data obtained indicate a high degree of interspecific differentiation of representatives of the genus Eremurus within the Almaty region, which confirms the provisions of the “Flora of Kazakhstan” and modern ideas about the morphological plasticity of the genus. The set of features, including habitus, size and shape of leaves, architecture of the inflorescence, characteristics of the perianth and an integral indicator of fruit size, reflects the adaptation of species to various ecological conditions of the region. Visual materials (Figures 4 to 9) combined with quantitative data (Table 2) make it possible to trace stable morphological differences between species that are not limited to intraspecific variability.
The height of plants varies in a wide range - from tall forms of Eremurus robustus (100-200 cm) to low-growing and compact species such as E. anisopterus (30-70 cm) and E. cristatus (50-70 cm). The results of the univariate analysis of variance (ANOVA) indicate a statistically significant effect of the “species” factor on plant height, which emphasizes the species-specific conditionality of habitus. The height of E. robustus combines with a powerful stem and large leaves (Figure 4) and can be interpreted as a sign of adaptation to more mesophytic or moderately humid conditions, whereas the decrease in height in E. anisopterus and E. cristatus reflects a xeromorphic strategy aimed at reducing transpiration losses and mechanical stress in arid and open habitats (Figures 7 and 9).
The width and shape of the leaves demonstrate one of the most distinct trends in morphological differentiation. The maximum leaf widths are typical for E. robustus (3.0-8.0 cm), whereas in E. tianschanicus, E. anisopterus and E. inderiensis, the leaves are narrowly linear or keeled (0.4-1.2 cm). ANOVA revealed significant interspecific differences in this trait, which confirms its high diagnostic and ecological significance. The narrowing of the leaf blade, grooving and keeling, recorded in a number of species (Figures 5, 8 and 9), can be considered as adaptive mechanisms aimed at reducing the area of the evaporating surface and increasing the efficiency of the water balance in conditions of increased insolation and moisture deficiency.
The architecture of the inflorescence also exhibits significant variability, ranging from narrowly cylindrical and dense inflorescences in E. tianschanicus, E. fuscus, and E. inderiensis to a broad and spreading form in E. anisopterus (Figures 5, 6, 8 and 9). These differences reflect specific features of generative organ spatial organization and are likely associated with variations in reproductive strategies and pollination conditions. Compact and dense inflorescences may offer greater resilience to wind load and temperature fluctuations, whereas spreading forms ensure wider exposure of flowers to pollinators in open habitats.
The size, shape, and color of the perianth demonstrate consistent interspecific variation. Perianth length varies from 8-10 mm in E. cristatus to 15-17 mm in E. robustus and reaches 20 mm in E. anisopterus (Table 2; Figures 4, 7 and 9). ANOVA confirmed statistically significant differences for this trait. Variations in perianth shape (rotate, narrowly campanulate, campanulate) and color (pale pink, yellow, tawny, purple hues with characteristic stripes) can be considered as elements of a species-specific reproductive complex, influencing interactions with pollinators and enhancing species isolation.
The ratio of stamen length to perianth length is a stable morphological characteristic that clearly distinguishes species. In E. tianschanicus and E. fuscus, the stamens exceed the perianth; in E. cristatus, the stamen filaments are 1.5 times longer than the perianth, whereas in E. robustus, they are somewhat shorter, and in E. anisopterus, they are significantly shorter than the corolla (Table 1; Figures 4 to 9). This trait possesses high taxonomic value and may be associated with the specific positioning of reproductive organs relative to the pollinator’s body.
The integral indicator - fruit diameter (M ± SD) - demonstrated the most pronounced interspecific variability. Minimal values were observed in E. fuscus and E. cristatus, whereas in E. robustus they were significantly higher, and maximum values were recorded in E. inderiensis and E. anisopterus (Table 2). ANOVA results showed a statistically significant influence of the “species” factor on fruit diameter, underscoring its key role in the reproductive differentiation of species. Larger fruits in E. inderiensis and E. anisopterus can be considered an adaptation ensuring increased seed viability under arid and unstable habitat conditions.
The combination of morphological data and the results of the analysis of variance confirm that the species of the genus Eremurus in the Almaty region form a clearly differentiated complex of morphological adaptations reflecting the ecological heterogeneity of the region. The most informative traits, statistically confirmed by ANOVA, are plant height, leaf width, perianth length, and fruit diameter. These traits possess high diagnostic value and can be recommended for comparative eco-morphological studies, as well as for refining the taxonomic status and adaptive strategies of Eremurus species.
From a taxonomic point of view, the most informative morphological traits for distinguishing the studied Eremurus species are plant height, leaf width and shape, inflorescence architecture, perianth length and shape, the position of stamens relative to the perianth, and fruit diameter. These characters showed clear interspecific differentiation and may be used as additional diagnostic features in comparative taxonomic studies of the genus. In particular, the compact habitus and narrow keeled leaves of E. anisopterus, the large fruit diameter of E. anisopterus and E. inderiensis, the robust habitus and broad leaves of E. robustus, and the smaller habitus with narrow leaves in E. cristatus provide useful morphological markers for species delimitation. Anatomical traits, including epidermal thickness, palisade mesophyll development, and the size of water-storage cells, also complement external morphological features and may strengthen species-level comparisons, especially among taxa occurring in ecologically contrasting habitats. However, these anatomical characters should be interpreted as supportive rather than independent taxonomic criteria and require further validation using broader population sampling and molecular data.
4.2. Anatomical adaptations of Eremurus species
The anatomical structure of leaves and stems in the studied Eremurus species demonstrates a consistent set of xeromorphic traits typical of plants adapted to arid and semi-arid environments. A single-layered epidermis with a well-developed cuticle and predominantly sunken stomata was observed in all investigated species. Such structural features are widely reported for xerophytic plants and are generally associated with reduced transpiration and increased resistance to water deficit under conditions of high solar radiation and strong temperature fluctuations.
Despite these shared xeromorphic characteristics, the studied species exhibit clear interspecific differentiation in quantitative anatomical traits. The variation observed in epidermal thickness, palisade mesophyll development, and the size of water-storage cells suggests that species of the genus Eremurus employ different structural strategies to cope with environmental stress.
The palisade mesophyll showed the greatest variation among species. Species occurring in sandy and highly arid habitats, such as E. anisopterus and E. inderiensis, exhibited a more developed palisade layer. This pattern is consistent with xeromorphic adaptation, as an increased palisade layer is often associated with plants growing under strong illumination and limited water availability. In contrast, species associated with rocky mountain slopes (E. cristatus, E. fuscus) showed comparatively moderate development of the palisade mesophyll, which may reflect differences in ecological conditions and light regimes.
Another important anatomical feature is the presence of well-developed water-storage cells in both leaf mesophyll and stem tissues. The diameter of these cells varied considerably among species and was particularly pronounced in E. anisopterus. The development of large water-storage cells in species associated with sandy desert habitats may reflect structural adaptation to irregular water availability, allowing temporary accumulation of moisture during favorable periods.
Stem anatomy also revealed patterns of interspecific differentiation. Species from arid environments, especially E. anisopterus, showed relatively thicker epidermal tissues and larger cortical parenchyma cells. These features may contribute to improved mechanical stability and water retention under conditions of mobile sandy substrates and intense solar radiation.
The results of ANOVA confirm that several anatomical traits-particularly palisade mesophyll thickness, epidermal thickness, and the diameter of water-storage cells-play a significant role in distinguishing the studied species. These traits therefore represent informative structural parameters reflecting adaptive differentiation within the genus.
The obtained results indicate that species of Eremurus demonstrate a combination of shared xeromorphic features and species-specific anatomical modifications. Such structural variability likely reflects the ecological heterogeneity of habitats in southeastern Kazakhstan, ranging from foothill steppe landscapes to sandy desert environments. The present findings highlight the importance of quantitative anatomical traits for understanding adaptive differentiation and ecological specialization within the genus.
These patterns are consistent with xeromorphic strategies commonly reported for plants in arid environments; however, in the absence of physiological or environmental measurements, they should be interpreted as structural correlations rather than direct evidence of adaptive mechanisms.
Although differences in anatomical traits were quantified and explored statistically, these results should be interpreted as indicative patterns rather than definitive adaptive differentiation.
The studied traits provide useful comparative descriptors, but their taxonomic value requires further validation using multivariate or integrative approaches.
5. Conclusions
The results demonstrate consistent interspecific differences in morphological and anatomical traits among the studied species; however, these patterns should be interpreted as indicative of structural differentiation under local environmental conditions rather than definitive adaptive or taxonomic relationships. Morphological parameters such as plant height, leaf width and shape, inflorescence structure, perianth size, and fruit diameter demonstrate consistent interspecific differences, statistically supported by ANOVA results, indicating their diagnostic and ecological significance. The anatomical structure of the leaf and stem is characterized by a common xeromorphic organization, including a single-layered epidermis with a developed cuticle, predominantly sunken stomata, differentiated mesophyll, and the presence of water-storage cells. Quantitative differences in palisade mesophyll thickness, epidermal structure, and the diameter of water-storage tissues reflect species-specific adaptive strategies to high insolation and limited water availability. Species associated with highly arid habitats, particularly Eremurus anisopterus and E. inderiensis, exhibit more pronounced xeromorphic traits, including compact habitus, narrow leaves, and well-developed water-storage tissues. In contrast, species occurring in foothill and mountain-steppe ecosystems (E. robustus, E. tianschanicus, E. fuscus, and E. cristatus) demonstrate a broader combination of morpho-anatomical characteristics, reflecting greater ecological plasticity. However, the present study has several limitations. The anatomical and morphological analyses were based on material collected during a single field season (2025) and from a limited number of populations. Therefore, the observed variability primarily reflects interspecific differentiation rather than full population-level variation. The observed patterns primarily reflect interspecific differentiation under local environmental conditions rather than complete species-wide variability. Future research should expand the sampling to multiple populations across the distribution range of each species and integrate additional approaches, including physiological and molecular analyses. Such studies would allow a more comprehensive understanding of the adaptive mechanisms and evolutionary differentiation of Eremurus species in arid and mountain ecosystems of Central Asia.
Data Availability Statement
The entire data set that supports the results of this study was published in the article itself.
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Edited by
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Editor:
Takako Matsumura Tundisi
