Habitat-driven variability in morphological and anatomical traits of Dactyloctenium aegyptium (Poaceae) in semi-arid regions

Abstract

Background

The ability of Dactyloctenium aegyptium (L.) Willd. to adapt to diverse habitats reflects its remarkable ecological plasticity.

Methods

This study investigates the morphological and anatomical traits of D. aegyptium across six distinct habitat types in semi-arid regions, which vary in moisture availability, soil texture, and levels of disturbance. We used one-way ANOVA and Principal Component Analysis (PCA) to evaluate habitat-driven differences and identify key patterns of trait variation.

Results

We found significant habitat-induced differences in morphological traits such as stem length, root length, leaf length, and plant biomass. Stem length and diameter peaked in sandy and canal bank habitats, while abandoned land exhibited the lowest values. Leaf length was highest in canal bank habitats and lowest in abandoned land. Sandy places supported the highest biomass, reflecting optimal growth conditions, while abandoned land recorded minimal biomass. Anatomical traits displayed habitat-specific adaptations, with roadside habitats exhibiting the thickest root epidermis and sandy places showing the highest endodermis thickness. Leaf anatomical features such as vascular bundle dimensions and bulliform cell density varied significantly, reflecting adaptations to environmental stresses. PCA revealed that plant traits clustered according to habitat type, with traits such as stem length, leaf size, and biomass associated with resource-rich habitats like roadside and agricultural land, indicating habitat-driven differentiation and adaptive plasticity.

Conclusion

These findings highlight the strong relationship between habitat variability and morphological and anatomical plasticity in D. aegyptium, providing insights into its adaptive strategies in semi-arid regions. This study contributes to understanding the ecological resilience of grass species under varying environmental conditions and highlights the importance of habitat-driven plasticity in shaping plant traits.

Supplementary Information

The online version contains supplementary material available at 10.1186/s12870-025-07361-5.

Introduction

Grass species are among the most widely distributed plant taxa, showcasing extensive variation in functional and phenotypic traits [1, 2]. These variations often arise due to environmental pressures, such as changes in geographic location, climate, soil conditions, and interactions with biotic and abiotic factors [3]. Over time, these selective pressures have driven the evolution of ecotypes, which are distinct populations within a species adapted to specific environmental conditions [4, 5]. Ecotypes provide an exceptional model for studying adaptive plasticity and the mechanisms underlying plant resilience in diverse habitats [6, 7]. Within the Poaceae family, anatomical traits exhibit significant variability, surpassing that of most other plant families [8]. These traits, including leaf thickness, stomatal density, sclerenchyma reinforcement, vascular bundle size, and bulliform cell distribution, are shaped by environmental stresses such as drought, salinity, and temperature fluctuations [9, 10]. These anatomical features not only provide insights into the taxonomic classification of grasses but also serve as indicators of environmental stress tolerance [11]. Structural features of vascular bundles—such as their size, the diameter of xylem vessels, and the arrangement of bundle sheath cells, play a crucial role in maintaining efficient water and nutrient transport under adverse environmental conditions [12, 13].

Dactyloctenium aegyptium (L.) Willd. is an annual grass species belongs to Poaceae family commonly known as Egyptian crowfoot grass. It is the fifth largest family of flowering plants consisting 50 tribes, 660 genera and near about 10,000 species. In Pakistan, Poaceae family comprising approximately 158 genera and 490 species. Some plants of this family used as medicine such as for the diseases like antidiabetic, antiulcer, antioxidant, diuretic and hypertension [14]. This species is known for its remarkable adaptability to a wide range of habitats, including temperate, tropical, and subtropical regions, as well as saline and semi-arid environments [15]. It thrives in regions of Africa, Asia, and the Middle East, often colonizing roadside areas, agricultural lands, and abandoned fields [16]. Anatomical adaptations, such as fibrous roots for water absorption, large cortical parenchyma cells for enhanced photosynthesis, and increased xylem and phloem diameters with enlarged vascular bundles, contribute to efficient transport of water and nutrients, enhancing resilience in challenging environments such as those with limited sunlight and low rainfall [17]. Morphologically, D. aegyptium exhibits features such as mat-like growth which is adapted to the habitats as wet places, which further enhance its ecological success by producing more seeds [18].

Despite its ecological importance, limited studies have explored the relationship between habitat variability and the morphological and anatomical traits of D. aegyptium. In Pakistan little is known about the ecological factors, particularly in the semiarid region of Punjab. In semi-arid regions, reduced rainfall can limit the growth of D. aegyptium, prompting anatomical adaptations such as increased parenchyma development for water storage [3]. This grass serves as an excellent model for investigating the role of habitat-driven plasticity in shaping plant form and function [19]. To address the knowledge gap regarding the ecological adaptability of D. aegyptium, this study investigates how morpho-anatomical traits vary across habitats that differ in environmental stressors and anthropogenic disturbance. The traits selected in this study were chosen based on their established ecological relevance to drought tolerance, water transport, and structural adaptation under semi-arid conditions [16]. While widely used traits such as SLA are important indicators of photosynthetic efficiency, our study focused on traits that could be measured consistently in field-based settings with limited resources, and which directly capture morpho-anatomical strategies of resilience. These habitats provide a natural experiment for exploring the species’ adaptive strategies. We particularly focus on morphological traits and anatomical traits which serve as indicators of structural and functional adaptations to specific ecological contexts. This approach allows us to better understand how D. aegyptium maintains resilience and functional performance across varied semi-arid landscapes.

Methodology

Study area

The semi-arid regions of Punjab, Pakistan, particularly in the vicinity of Okara and Kasur located at the southeast side of Lahore (31°12′ N and 74°44′ E) are characterized by unique geographical and ecological features (Fig. 1). These areas consist of predominantly flat to gently undulating terrain, interspersed with extensive irrigation networks that underpin agricultural productivity [2]. The climate is distinctly semi-arid, with hot summers and mild winters. Mean annual Rainfall is 627.12 mmyr − 1 ± 205.33 mmyr − 1 during the time period from1981-2020, and primarily occurs during the monsoon season [19]. The mean annual temperature ranges from 23 to 26 °C, with a 16–19 °C T_min and of 29–33 °C T_max. Fertile alluvial soils dominate the irrigated plains, providing an ideal foundation for intensive agriculture, including the cultivation of staple crops like wheat and rice, alongside a variety of vegetables [20]. The vegetation is a dynamic mix of native species adapted to semi-arid conditions and cultivated crops, creating a landscape that is both ecologically diverse and agriculturally productive [21]. The extensive irrigation infrastructure plays a pivotal role in sustaining agriculture, effectively mitigating the challenges posed by water scarcity inherent to semi-arid climates.

Fig. 1.

Study area map showing the location of various habitats for D. aegyptium sampling in semi-arid region

Sample collection

In the semi-arid region of Punjab, six habitats were selected for the study, including roadside, canal banks, agriculture land, moist places, abandoned land, and sandy places. The six habitats selected in this study represent distinct ecological conditions shaped by a combination of environmental variability and anthropogenic influence (Supplementary Table 1). Roadside habitats are typically exposed and compacted, with poor soil structure, high levels of disturbance from vehicular activity, and fluctuating moisture availability. Canal banks, in contrast, are semi-natural zones adjacent to irrigation channels, characterized by periodic waterlogging, loamy soils, and relatively low levels of disturbance. Agricultural lands are intensively managed environments where regular tillage, irrigation, and fertilizer application create moderately fertile soils with intermediate disturbance. Moist places are low-lying microhabitats that maintain stable moisture levels due to surface runoff or subsurface seepage, often supporting higher levels of organic matter accumulation. Sandy places, defined by their loose, well-drained soils, are nutrient-poor and retain little moisture, presenting conditions of high edaphic stress. Abandoned lands are previously cultivated or disturbed sites that have been left unmanaged, typically exhibiting signs of ecological degradation, low soil fertility, and minimal vegetative cover. These habitats were purposefully selected to capture a broad range of environmental stressors and resource availabilities, thereby enabling a detailed assessment of trait plasticity and ecological adaptation in D. aegyptium across gradients of moisture, disturbance, and soil condition. In each habitat type, three independent sampling locations were established to capture spatial variability. From each location, two mature and healthy individuals of D. aegyptium were collected at spatial intervals of approximately 5–10 m, ensuring that plants were not clustered and reducing microhabitat bias. This design resulted in six individuals per habitat and a total of 36 plants across all six habitats. For anatomical analyses, one standardized root, stem, and leaf segment was taken from each plant, yielding 108 organ samples. To maintain consistency, only individuals from areas where D. aegyptium was the dominant species, with minimal overlap from other grasses, were selected. While community-level metrics such as total coverage, species density, and vegetation height were not formally recorded in this study, all selected sites reflected typical habitat conditions for this species within the semi-arid landscape. The collected plants were identified by Dr. Fahim Arshad and have been deposited (Deposit number NM 01) in the Herbarium of the Department of Botany, University of Okara. After morphological measurements, plant samples were preserved for anatomical analysis. Root, stem, and leaf segments, each approximately 3.81 cm in length for dipping in solution, were cut and stored in a 70% methanol solution to maintain tissue integrity.

Morphology traits measurement

For the fresh weight measurement, the plant samples were collected and were taken to the laboratory immediately, and the weight of the plant in the analytical balance [22, 23]. Height calculated from base to tip of largest branch. A measuring scale was used to measure the plant height in cm [22]. Number of nodes and internodes was counted at the time of sample collection. Stem diameter was measured by using a Vernier caliper from the specified point such as 10 cm above the soil surface. The root length is measured by measuring scale and its diameter was measured by Vernier calipers [22].

Anatomy traits measurement

For anatomical analysis, one healthy root, stem, and leaf segment (each approximately 3.8 cm in length) was excised from every specimen using sterilized stainless-steel blades (Supplementary Table 2). The samples were immediately fixed in 70% ethanol for 24 h to preserve tissue integrity and cellular structure. As the plant material was sufficiently soft, no dehydration or embedding was required, and freehand sections were prepared without microtomy. Thin transverse sections were obtained manually using a sharp razor blade. The prepared sections were first stained in a 1% safranin solution (1 g safranin in 100 mL distilled water) for 2–5 min to highlight Lignified tissues, then gently rinsed with distilled water. Counterstaining was performed using 1% fast green solution prepared in 95% ethanol, allowing contrast between different tissue types. Excess stain was blotted with tissue paper, and the sections were mounted in glycerin on clean glass slides. Coverslips were carefully applied and sealed to prepare the slides for microscopic examination. Prepared slides were examined under a compound Light microscope at magnifications of 4×, 10×, and 40× [24]. Anatomical features were observed and recorded for each organ type. In the root, parameters measured included rhizodermis thickness, cortex cell area, pith cell area, xylem area, phloem area, and endodermis thickness. In the stem, measurements included epidermis and hypodermis thickness, cortex area, xylem and phloem area, vascular bundle length and width, and sclerenchyma thickness. In the leaf, observations covered adaxial and abaxial epidermis thickness, mesophyll cell area and thickness, xylem and phloem area, vascular bundle dimensions, bulliform cell presence, and stomatal area.

Data analysis

Statistical analyses were performed to evaluate habitat-based differences in morphological and anatomical traits of D. aegyptium. One-way analysis of variance (ANOVA) was used to test for significant differences among habitat groups, followed by Tukey’s HSD post-hoc test for pairwise comparisons. Mean and standard error (SE) were calculated for each trait, and results were reported as mean ± SE with corresponding p-values. Principal Component Analysis (PCA) was performed to identify the major axes of variation and visualize habitat differentiation based on morphological and anatomical traits. Data were standardized prior to PCA using scikit-learn [25], and biplots were generated to show variable contributions to PC1 and PC2 and habitat clustering. All analyses and visualizations were conducted in Python 3.11 using scipy.stats, scikit-learn, pandas, matplotlib, and seaborn [26].

Results

Morphological traits variability

Morphological traits of D. aegyptium exhibited significant habitat-driven variability, reflecting its adaptive plasticity. Stem length and diameter were highest in sandy (46.13 ± 0.96 cm; 1.47 ± 0.15 cm) and moist habitats, while abandoned land showed the lowest values (30.17 ± 1.61 cm; 0.75 ± 0.15 cm; p < 0.05) (Supplementary table). Leaf length varied significantly (p < 0.001), peaking in canal bank habitats (43.90 ± 1.39 cm) and declining in abandoned land (26.77 ± 1.90 cm). Root length was longest in agriculture land (5.17 ± 0.19 cm; p < 0.01) and shortest in canal bank (3.31 ± 0.21 cm). Sandy places supported the highest node count (8.41 ± 0.17), fresh weight (8.66 ± 0.34 g), dry weight (1.56 ± 0.17 g), and plant biomass (10.24 ± 0.55 g), while abandoned land recorded the lowest values (p < 0.001). The PCA analysis for morphological traits demonstrated that the first principal component (PC1) explained 50.4% of the variation, while the second principal component (PC2) accounted for 19.8% (Fig. 2a). Traits such as the number of nodes, fresh weight, and root length contributed significantly to the variation observed in PC1, while traits such as stem length and dry weight influenced PC2. The distribution of habitat groups showed clear clustering, root length strongly associated with plants in agricultural and abandoned lands. Conversely, plants from roadside and sandy habitats exhibited greater leaf width and stem diameter, as indicated by their higher loading on the PCA axes.

Fig. 2.

Principal Component Analysis biplots showing trait variation in D. aegyptium across six different habitats. Each subplot represents a specific trait category: (a) morphological traits, (b) root anatomical traits, (c) stem anatomical traits, and (d) leaf anatomical traits. The axes (PC1 and PC2) represent the first and second principal components, indicating the percentage of total variance explained. Vectors represent individual trait contributions to overall variation, with longer arrows indicating higher influence. The color intensity of arrows corresponds to trait contribution magnitude. Colored ellipses and points denote habitat groupings: red (abandoned land), cyan (agriculture land), green (canal bank), magenta (moist places), pink (roadside), and black (sandy places). Traits are labeled along vectors for reference, and clustering reflects trait–habitat associations

Anatomical traits variability

Root traits of D. aegyptium demonstrated significant habitat-specific variability, emphasizing its adaptability to contrasting ecological conditions (Fig. 3). Epidermis thickness varied significantly (p < 0.01), peaking in roadside (14.96 ± 1.34 μm) and sandy places (13.55 ± 1.31 μm), while abandoned land had the lowest value (7.28 ± 0.72 μm). Endodermis thickness was highest in sandy places (11.55 ± 1.30 μm; p < 0.001) and lowest in agriculture land (4.24 ± 0.73 μm). Root cortex thickness (p < 0.001) peaked in moist places (98.83 ± 3.74 μm) and was thinnest in abandoned land (54.10 ± 2.10 μm) (Supplementary Table3). Root xylem and phloem diameters also varied significantly (p < 0.001), with roadside habitats exhibiting the largest diameters (55.58 ± 1.86 μm and 64.16 ± 1.61 μm, respectively) and abandoned land the smallest (27.69 ± 2.57 μm and 19.83 ± 1.46 μm). Root pith diameter was largest in roadside habitats (247.61 ± 5.16 μm) and smallest in abandoned land (97.08 ± 3.37 μm). For root traits, PC1 and PC2 captured 58.8% and 17.7% of the total variation, respectively (Fig. 2b). Root cortex and root xylem were the most influential contributors to PC1, while epidermis and endodermis traits significantly impacted PC2. Plants from moist places and canal banks strongly associated with root cortex and epidermis. In contrast, sandy places and roadsides were characterized by root xylem and endodermis traits.

Fig. 3.

Transverse sections showing root anatomy of D. aegyptium collected from six different habitats under light microscopy. Images display the structural organization of vascular and ground tissues, highlighting variations in root diameter, cortical thickness, and vascular bundle arrangement. (a) Roadside; (b) Canal bank; (c) Moist places; (d) Agricultural land; (e) Sandy places; (f) Abandoned land

Stem traits of D. aegyptium showed significant habitat-driven variability, reflecting structural adaptations to environmental conditions (Fig. 4). Epidermis thickness (p < 0.05) was highest in moist places (14.75 ± 0.60 μm) and lowest in agriculture land (7.08 ± 0.90 μm) (Supplementary Table 3). Hypodermis thickness varied markedly (p < 0.001), peaking in moist places (168.81 ± 7.04 μm) and sandy places (148.74 ± 5.16 μm), while abandoned land recorded the lowest value (39.30 ± 3.17 μm). Xylem (p < 0.01) and phloem diameters (p < 0.001) were largest in sandy places (60.74 ± 5.98 μm and 60.17 ± 5.32 μm, respectively) and smallest in abandoned land (28.06 ± 5.64 μm and 26.66 ± 3.87 μm). Vascular bundle (VB) length and width also varied significantly (p < 0.01), with moist places (175.92 ± 4.45 μm and 213.00 ± 4.22 μm, respectively) and sandy places (162.12 ± 3.81 μm and 214.83 ± 5.01 μm) having the largest bundles, while agriculture land and abandoned land recorded the smallest values. Sclerenchyma thickness (p < 0.001) was highest in canal bank (25.81 ± 3.38 μm) and roadside habitats (24.82 ± 2.48 μm) and lowest in abandoned land (13.11 ± 1.16 μm), reflecting structural reinforcement in favorable environments. The PCA for stem traits revealed that PC1 accounted for 70.3% of the variation, with PC2 contributing 11.4% (Fig. 3c). Traits such as sclerenchyma and vascular bundle width were the dominant variables contributing to PC1, whereas phloem and vascular bundle length influenced PC2. Plants in agricultural lands and canal banks were associated with traits like sclerenchyma, while plants in sandy places and roadside habitats showed a stronger correlation with phloem and vascular bundle length.

Fig. 4.

Transverse sections of stem tissue in D. aegyptium collected from six distinct habitats, highlighting variation in anatomical features across environments. Sections reveal the epidermis, hypodermis, cortex, and arrangement of vascular bundles. (a) Roadside; (b) Canal bank; (c) Moist places; (d) Agriculture land; (e) Sandy places; (f) Abandoned land. Images were taken under light microscopy

Leaf traits of D. aegyptium exhibited significant habitat-driven variability, reflecting adaptive plasticity (Fig. 5). Adaxial epidermis thickness (p < 0.001) was highest in canal bank habitats (23.28 ± 1.26 μm) and sandy places (18.07 ± 1.46 μm), while abandoned land recorded the thinnest epidermis (10.55 ± 1.37 μm) (Supplementary Table3). Abaxial epidermis thickness (p < 0.01) peaked in moist places (17.20 ± 0.68 μm) and was lowest in roadside (9.14 ± 0.71 μm). Leaf cortex dimensions varied significantly (p < 0.001), with the largest cells observed in moist places (428.49 ± 28.92 μm) and agriculture land (411.52 ± 26.24 μm), while roadside habitats showed the smallest (209.30 ± 7.33 μm). Vascular bundle (VB) dimensions also differed, with moist places having the widest VB width (173.96 ± 5.04 μm; p < 0.001) and sandy places the longest VB length (71.99 ± 5.61 μm), while abandoned land exhibited the smallest VB width (40.42 ± 3.07 μm) and length (36.75 ± 3.37 μm). Xylem and phloem thickness (p < 0.001) were highest in moist places (51.50 ± 2.06 μm) and agriculture land (48.54 ± 3.38 μm), respectively, and lowest in abandoned land (25.17 ± 1.69 μm and 15.65 ± 2.12 μm). Bulliform cell density (p < 0.001) peaked in sandy places (164.03 ± 6.90 μm) and was lowest in moist places (128.25 ± 1.81 μm). Leaf traits showed that PC1 and PC2 accounted for 39.9% and 25.8% of the variation, respectively (Fig. 3d). Leaf cortex and bulliform cells were prominent contributors to PC1, while traits such as leaf vascular bundle length and adaxial epidermis were significant for PC2. Plants collected from agricultural lands and canal banks exhibited greater sclerenchyma thickness, while those from sandy places and roadside habitats showed increased phloem area and vascular bundle length. These trait patterns reflect habitat-specific structural adaptations captured in the PCA analysis.

Fig. 5.

Transverse sections of D. aegyptium leaf blades from six habitats, illustrating anatomical variation in epidermis, mesophyll, and vascular structures under light microscopy. (a) Roadside; (b) Canal bank; (c) Moist places; (d) Agriculture land; (e) Sandy places; (f) Abandoned land

Association between traits

The correlation analysis revealed that plant biomass was strongly positively correlated with stem length, leaf length, and fresh weight, highlighting their key contributions to growth. Fresh weight also correlated strongly with leaf length and moderately with stem length. Stem and leaf traits showed strong positive relationships, indicating coordinated growth. Weak positive correlations were observed between stem diameter and dry weight, as well as between the number of nodes and fresh weight (Fig. 6a). Negative correlations were weak, with root length showing minor trade-offs with aboveground traits. Strong positive correlations were observed between root phloem, root pith, and root xylem, highlighting their coordinated role in vascular transport and support structures. Epidermis thickness showed a strong positive correlation with root xylem and root pith, suggesting its association with structural adaptations. Moderate positive correlations were identified between the endodermis and root pith, reflecting shared roles in water and nutrient regulation. Root cortex exhibited weak correlations with other traits, emphasizing its relatively independent role in root anatomy (Fig. 6b). Strong positive correlations were observed between vascular bundle (VB) width and VB length, as well as between VB width and xylem thickness, highlighting their role in vascular transport efficiency. Phloem thickness exhibited a strong positive correlation with hypodermis and sclerenchyma, reflecting its contribution to translocation. Hypodermis thickness was moderately correlated with epidermis and VB dimensions, indicating its structural importance. Epidermis and sclerenchyma showed weaker positive correlations with most traits, suggesting their specialized roles (Fig. 6c). Strong positive correlations were observed between vascular bundle width and abaxial epidermis, as well as between VB width and VB length, highlighting the interdependence of vascular dimensions. Moderate positive correlations were identified between VB length and leaf cortex cell size, reflecting the role of bundle dimensions in supporting leaf structure. Leaf xylem showed weak positive correlations with epidermal traits and VB dimensions, emphasizing its role in water transport. In contrast, leaf phloem exhibited weak positive correlations with bulliform cells, reflecting a potential relationship between nutrient transport and leaf folding mechanisms (Fig. 6d).

Fig. 6.

Correlation matrices representing the relationships between morphological and anatomical traits of D. aegyptium across six semi-arid habitats. Each matrix displays Pearson correlation coefficients, where values range from − 1 to + 1 and are color-coded from purple (negative correlation) to yellow (positive correlation). (a) Morphological traits; (b) Root anatomical traits; (c) Stem anatomical traits; (d) Leaf anatomical traits

Discussion

The present study provides comprehensive insights into the morphological and anatomical adaptations of D. aegyptium, demonstrating its capacity to thrive across diverse habitats in semi-arid regions. These findings emphasize the intricate interaction between microhabitat and plant trait variability, revealing the adaptive strategies employed by this species to overcome ecological constraints.

Morphological adaptations: balancing growth and resource allocation

The significant variability in morphological traits across habitats reflects the ability of D. aegyptium to allocate resources strategically based on habitat conditions. Longer shoot lengths observed in habitats, such as roadside and sandy places, align with strategies for optimizing photosynthesis and minimizing competition for light under limited moisture conditions [27]. This result is consistent with studies by Palta et al. [28], who noted increased shoot elongation in dry environments to enhance competitive advantage. Conversely, shorter shoot lengths in abandoned lands and agriculture sites highlight constraints imposed by nutrient availability and competition [29]. Root length, another critical adaptive trait, was significantly reduced in water-saturated soils, such as canal bank habitats. This reduction is likely due to low oxygen availability in saturated soils, which can limit aeration and inhibit root elongation in wet conditions. Longer root systems in roadside and sandy habitats, where soil moisture is limited, suggest enhanced investment in belowground structures to access deeper water reserves [30]. Leaf morphology exhibited a clear response to habitat moisture gradients. Longer leaves in moist and canal bank habitats, reflect greater water availability and reduced need for structural conservation [31]. In contrast, shorter leaves in abandoned and roadside habitats minimize water loss through transpiration, showcasing the species’ plasticity. Interestingly, leaf width was also larger in moist places and canal bank habitats, suggesting enhanced photosynthetic capacity in resource-rich conditions [32]. These morphological adjustments illustrate the species’ ability to prioritize growth or conservation based on environmental constraints.

Anatomical traits: structural and functional adaptations

The anatomical traits of D. aegyptium provide a deeper understanding of its physiological responses to habitat variability. Root anatomical features, such as thicker Rhizodermis and endodermis in sandy and roadside habitats, highlight their critical role in reducing water loss and improving water absorption under drought stress [33]. This finding is consistent with Smith et al. [34], who emphasized the importance of epidermal adaptations in water-limited environments. Larger cortical areas in moist places and canal banks enhance storage capacity, buffering against temporary water deficits [14]. These results support the idea that cortical plasticity is a key adaptation for plants in fluctuating environments [35]. The vascular structures of the root, including xylem and phloem dimensions, exhibited notable habitat-driven variations. Larger xylem vessels observed in sandy places and roadside habitats improve water transport efficiency, supporting adaptation to limited moisture conditions [36], while thicker phloem in moist environments suggests an increased capacity for translocation and assimilation [37]. These adaptations reflect the dual demands of supporting growth and coping with stress, displaying the functional integration of root anatomy [38]. In stems, vascular bundle dimensions were significantly larger in sandy and moist habitats, suggesting enhanced hydraulic conductivity and structural support [39]. These traits are critical for maintaining growth under varying moisture conditions [40], as larger vascular bundles allow for efficient transport of water and nutrients [41]. Hypodermis and sclerenchyma thickness varied across habitats, providing mechanical stability and protection against environmental stressors such as wind or herbivory [42].

Leaf traits: photosynthetic efficiency and water conservation

Leaf anatomical traits, including vascular bundle size and bulliform cell density, exhibited clear habitat-specific patterns. Larger vascular bundles in canal bank habitats support greater hydraulic efficiency, facilitating sustained photosynthesis in water-rich conditions [43]. Conversely, the increased density of bulliform cells in sandy and roadside habitats reflects a water-saving mechanism, as these cells promote leaf folding during periods of water scarcity [44]. Thicker epidermal layers in moist habitats serve as a protective barrier against excessive transpiration, while thinner layers in drier areas minimize resource investment in non-photosynthetic tissues [45]. The contrasting sizes of xylem and phloem in leaf vascular bundles further reflect habitat-specific strategies. Larger xylem in drier habitats optimizes water transport, while thicker phloem in moist areas supports greater carbohydrate translocation, ensuring energy availability for growth [46]. These findings highlight the complementary roles of xylem and phloem in supporting ecological plasticity across environments.

Synergistic relationships between traits

The observed correlations between morphological and anatomical traits suggest a coordinated strategy to optimize growth and survival. Strong positive associations between shoot length, leaf length, and vascular bundle size emphasize the functional integration of aboveground and vascular systems. This synergy enables D. aegyptium to maximize resource acquisition and utilization, particularly in habitats with moderate moisture availability, such as canal banks and sandy soils. In water-scarce environments such as sandy places, correlation patterns suggest a potential trade-off between root traits (e.g., root length) and shoot-related traits (e.g., stem length, leaf length, and fresh weight), where enhanced investment in root systems may compensate for reduced aboveground growth. This allocation shift may reflect an adaptive response to limited water availability, consistent with findings from previous studies on drought-tolerant species [47–49].

Ecological implications and adaptive significance

The habitat-specific plasticity of D. aegyptium highlights its ecological resilience and adaptability to semi-arid environments. By modulating growth patterns and anatomical structures, the species effectively balances resource acquisition, water conservation, and mechanical stability. These adaptations enhance its survival across diverse habitats and likely contribute to its ecological success and widespread occurrence in semi-arid ecosystems, as reported in previous studies [15, 16]. This study contributes to our understanding of plant responses to environmental heterogeneity, emphasizing the importance of morphological and anatomical traits in shaping ecological success. The findings also have practical implications for land management, as D. aegyptium’s adaptability makes it a potential candidate for soil stabilization and erosion control in degraded landscapes. Future research should explore the genetic and physiological mechanisms underlying these adaptations, particularly in the context of climate change and increasing anthropogenic pressures. The morphological and anatomical plasticity of D. aegyptium highlights its remarkable ability to thrive in semi-arid regions. This resilience not only reflects its ecological versatility but also provides valuable insights into the strategies employed by plants to navigate complex environmental challenges.

Limitations

While this study provides valuable insights into the morphological and anatomical plasticity of D. aegyptium across diverse semi-arid habitats, it is important to acknowledge certain limitations. Environmental parameters such as soil moisture, nutrient content, and pH were not directly measured, and habitat classifications were based on field observations and existing ecological knowledge. Additionally, the sample size per habitat was relatively limited, which may constrain the generalizability of trait variation patterns. Future studies incorporating systematic soil analyses, larger sample sizes, and community-level assessments would enhance the ecological resolution and mechanistic understanding of habitat-driven plant adaptations.

Conclusion

This study explains the ecological adaptability of D. aegyptium in semi-arid regions through a comprehensive analysis of morphological and anatomical traits across varied habitats. Morphological traits such as stem length, leaf dimensions, and biomass demonstrated significant habitat-driven plasticity, reflecting the plant’s capability to optimize growth in resource-rich environments (e.g., moist places, canal banks, agricultural land) as well as under resource-limited conditions (e.g., sandy places, roadside, and abandoned land). Notably, stem and leaf dimensions were maximized in habitats with higher soil moisture, such as canal banks, aligning with previous findings on moisture-facilitated elongation. Conversely, traits like root length and epidermis thickness were pronounced in drier environments, such as sandy and roadside habitats. Anatomical investigations further revealed habitat-specific modifications, particularly in root and stem vascular structures, which enhanced resource transport efficiency. For instance, increased vascular bundle dimensions in moist habitats supported enhanced physiological activity, whereas thicker root cortex and epidermis in arid zones highlighted structural adaptations for water retention. The presence of bulliform cells in leaves from sandy habitats emphasizes their role in reducing transpiration under water stress, a crucial survival mechanism in arid climates. These findings collectively highlight the remarkable plasticity of D. aegyptium, enabling it to thrive across diverse habitats with varying ecological pressures. This adaptability highlights the plant’s potential for ecological restoration and as a model species for understanding plant responses to environmental heterogeneity.

Authors’ contributions

Conceptualization, M.W. and N.M.; Methodology, M.W; software, M.W. and F.A.; validation, M.W. and F.A.; formal analysis, M.W., U.A., and N.H.; investigation, N.M.; resources, U.A. and F.A.; Data curation, S.U.; writing—original draft preparation, M.W., N.M; writing—review and editing, M.W., V.S., M.S., S.U., N.H., N.M., A.A., M.T., A.R.O., U.A, and F.A.; visualization, M.W.; supervision, M.W. and N.H.; project administration, F.A.; funding acquisition, A.A. All authors having substantial contributions in research, read and agreed to the published version of the manuscript.

Funding

The authors extend their appreciation to the Deanship of Research and Graduate Studies at King Khalid University for funding this work through Large Research Project under grant number RGP2/310/46.

Data availability

Data are contained within the article.

Declarations

Ethics approval and consent to participate

All necessary permissions for plant collection were secured before conducting the study. Sampling was carried out on both public and private land, with verbal permission obtained from the respective landowners.

Consent for publication

Not applicable—this manuscript has no personal data from the authors.

Competing interests

The authors declare no competing interests.