Influence of Rhizosphere Dynamics and Soil Chemical Properties in Arid Environments on the Distribution, Abundance, and Diversity of Arbuscular Mycorrhizal Fungi (AMF)

Saud S. Aloud; Fahad Alotaibi; Salah N. Sorrori; Basil Alshebe

doi:10.3390/ecologies6040080

,

and

¹

Soil Science Department, College of Food and Agriculture Sciences, King Saud University, Riyadh 13362, Saudi Arabia

²

Plant Production Department, College of Food and Agriculture Sciences, King Saud University, Riyadh 13362, Saudi Arabia

³

Department of Environmental Science, University of Arizona, 429 Shantz Bldg, Tucson, AZ 85721, USA

^*

Author to whom correspondence should be addressed.

Ecologies2025, 6(4), 80;https://doi.org/10.3390/ecologies6040080
(registering DOI)

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Abstract

This study investigates the influence of rhizosphere dynamics and soil chemical properties on the distribution, abundance, and diversity of arbuscular mycorrhizal fungi (AMF) across two seasons (summer and winter). A total of 11 rhizospheric soil and root samples were collected from various wild plant species, including Senna italica, Cyperus laevigatus, Phragmites australis, Pelargonium peltatum, Zygophyllum simplex, Citrullus colocynthis, Malva parviflora, Zygophyllum coccineum, Calotropis procera, Solanum nigrum, and Salsola baryosm. Phragmites australis exhibited the highest AMF spore count (175 and 124/100 g dry soil in summer and winter), while Calotropis procera showed the lowest (101 and 63/100 g). AMF species identified included Glomus ambisporum, Rhizophagus intraradices, Claroideoglomus etunicatum, Diversispora globifera, Funneliformis geosporum, Funneliformis mosseae, Rhizophagus fasciculatus, and Gigaspora spp. The Shannon diversity index ranged from 0.692 (Zygophyllum simplex) to 0.653 (Salsola imbricata), and Simpson’s index from 0.498 to 0.461. Phragmites australis recorded the highest root colonization (90.5% and 84.7%), arbuscule (76% and 69.3%), and vesicle formation (36%) in summer, while Calotropis procera had the lowest. In summer, AMF spore counts showed significant correlations with soil nutrients (N, P, K), and with total organic carbon (TOC) and organic matter (OM). During winter, TOC and OM remained influential, while correlations with nutrients weakened. Soil pH, electrical conductivity (EC), and texture exhibited minimal correlation with AMF spore counts in both seasons.

Keywords:

rhizosphere dynamics; soil chemical properties; wild plants; AMF

1. Introduction

Arbuscular mycorrhizal fungi (AMF) are globally widespread and constitute a significant component of the soil microbial community, forming symbiotic associations with nearly 80% of wild plants and numerous agricultural crops [,]. These fungi play a critical role in ecosystem functioning, influencing nutrient uptake, productivity, and providing plants with enhanced defense mechanisms against various environmental and agricultural stresses, including drought, salinity, and acidity. Despite their ecological importance, the distribution, abundance, and diversity of AMF are influenced by various environmental and soil factors, including root density, soil physicochemical properties, and seasonal changes [,,,]. Plant root density significantly affects AMF spore counts in the soil. Researchers such as Cuenca and Lovera and Anderson et al. have reported that AMF spores may be nearly absent in soils with diminished root densities. This indicates the crucial role that root presence plays in the distribution and abundance of AMF, as these fungi rely on host plants for their development and symbiotic functions. Additionally, Brundrett highlighted that AMF spore production is influenced by plant phenology, further emphasizing the interplay between plant developmental stages and AMF dynamics [,,].

Soil characteristics, including pH, moisture content, and disturbance such as high salinity, are important determinants of AMF distribution and abundance. Soil pH has been a focus of numerous studies, with some researchers reporting no significant correlation between AMF spore counts and pH levels due to changes in soil chemistry. However, Shukla et al. found a positive correlation between AMF spore populations and soil pH, indicating that soil pH may play a more nuanced role in influencing fungal populations [,,,,].

In addition to pH, soil organic matter content is a key factor in AMF distribution. Oehl et al. suggested that a decrease in organic content may reduce AMF spore numbers and presence in the soil. Moreover, soil disturbances, such as plowing and land use changes, have been shown to negatively impact AMF spore counts [,,].

Seasonal changes significantly influence AMF colonization and spore production. Titus et al. noted that seasonal changes affect AMF colonization by influencing the developmental and physiological stages of the host plant. Khan found that AMF root colonization and endogone spore abundance fluctuate seasonally, with spore numbers peaking during the months of October to January. Similarly, Guadarrama and Álvarez-Sánchez reported that AMF species diversity and spore counts are higher during the dry season compared to the rainy season, highlighting the impact of seasonal moisture variations on AMF communities. Studies by Adil et al. also indicated that seasonal variations in temperature and soil moisture have a marked effect on AMF spore production, and the abundance of AMF populations is significantly influenced by the season. This underscores the challenge of establishing clear patterns of AMF distribution and abundance due to seasonal fluctuations [,,,].

The influence of edaphic and climatic factors on AMF dynamics is well-documented. Lingfei et al. indicated that edaphoclimatic characteristics affect both the temporal and spatial dynamics of AMF infection and spore populations. Soil moisture, pH, and depth are key factors affecting AMF spore populations with increasing soil depth. Further studies have shown that soil fungal diversity tends to be higher in environments with lower pH levels. Climate factors, including temperature, precipitation, and cold tolerance, also play significant roles in AMF distribution. The interaction between soil fungi and their environment, including competition and mutualism, may further influence AMF community structure [,,,].

Recent molecular studies using high-throughput sequencing have deepened our understanding of AMF communities, particularly in relation to rhizosphere processes and soil chemistry. For example, Zhang et al. found that rhizosphere soils often harbor greater AMF diversity than plant roots, likely due to microenvironmental factors such as root exudates and localized nutrient availability. Likewise, Luo et al. reported strong correlations between soil properties—such as pH, organic matter, phosphorus, and nitrogen—and AMF richness across arid desert landscapes. Spatial heterogeneity, driven by topography, moisture gradients, and host plant identity, further contributes to variation in the abundance of AMF genera such as Glomus and Diversispora [,].

Despite this growing body of research, knowledge of AMF dynamics in arid and semi-arid environments remains limited—particularly regarding seasonal changes in colonization and diversity in wild plant communities. In regions such as Riyadh, where extreme climate conditions prevail, such knowledge is essential for understanding plant–fungal interactions and ecosystem resilience. Therefore, this study aims to investigate how selected physical and chemical soil properties affect AMF spore abundance, species richness, and root colonization intensity. By comparing samples collected in two distinct seasons, we assess how environmental variability shapes AMF communities in the wild ecosystems surrounding the Riyadh region.

2. Materials and Methods

2.1. Sampling Collection Strategies

Eleven species of wild plants associated with AMF were targeted to study the seasonal variation in AMF such as the following: Senna italica, Cyperus laevigatus, Phragmites australis, Pelargonium peltatum, Zygophyllum simplex, Citrullus colocynthis, Malva parviflora, Zygophyllum cocceinum, Calotropis procera, Solanum nigrum, Salsola baryosm, where all samples were collected during two different seasons. Both soil and plant roots samples were gathered from the rhizosphere zone of the targeted plants at a depth of 3–10 cm. Three replicates were randomly collected per plant during summer and winter seasons, respectively, to obtain a total of 33 samples per season. All samples were put into polythene bags and labeled with data of samples such as plant name, date, and location; then, we brought them to the laboratory. In lab, root samples were thoroughly cleaned with tap water to remove adhered soil, and fine roots were carefully separated from the whole mass of roots. Fine roots were cut into 1 cm fragments and immersed in FAA solution in glass tubes and stored at 4 °C for later analysis. The soil samples were air-dried at lab temperature for 15 days then stored at 4 °C for soil physicochemical properties analysis, AMF spore counts, abundance, and diversity [].

2.2. Extraction of AMF Spores

AMF spores were extracted according to wet sieving and decanting method by Khan, and AMF spores were counted as numbers/100 g soil; spores were identified using synoptic keys provided on the INVAM website and Schenck and Perez manual (1990) [].

2.3. Measurement AMF Colonization Ratio

To measure AMF colonization ratio in roots, fresh fine roots were carefully selected, stained, and studied according to methods of Phillips and Hayman (1970) and Al Qarawi et al. (2012) [,].

2.4. Soil Physicochemical Characteristics

Soil pH and electrical conductivity (EC) were assessed using a 1:1 mixture of soil and deionized water and measured by a pH meter and an EC meter. The distribution of soil particle sizes was examined using the hydrometer technique, and soil texture was classified according to the USDA soil textural triangle. Cation exchange capacity (CEC) was evaluated through extraction with 1 N ammonium acetate (NH₄OAc) at pH 7.0. Concentrations of exchangeable cations (Ca²⁺, Mg²⁺, K⁺, and Na⁺) were determined in the extract using inductively coupled plasma–optical emission spectrometry (ICP-OES). Calcium carbonate (CaCO₃) content was quantified using the calcimeter technique. Soil organic matter (OM) was estimated via the Walkley–Black wet oxidation method [,].

2.5. Statistical Analyses

The acquired data were arranged, and descriptive statistical analyses including mean, minimum, maximum, standard deviation (SD), and coefficient of variance (CV) were performed using SPSS 27 (IBM Corp., New York, NY, USA) and Microsoft Excel. The figures were drawn using Microsoft Excel program. Comparison of the mean was made with Statistical Package for Social Studies (SPSS 27; IBM Corp., New York, NY, USA) using least significant difference at the risk level of 5% [].

3. Results and Discussion

The average values of physiochemical properties of soils are given in Table 1, which indicates that the soil texture was a sandy loam with an alkaline pH of 8.1, moderate salinity (EC 2.8 dS/m), and high calcium CaCO₃ content of (11.91%) which is a typical of arid environments. These conditions can restrict nutrient availability and make it more difficult for AMF to colonize roots. For instance, sandy soils have low water retention, alkaline pH limits phosphorus availability, and salinity can inhibit fungal growth. Despite these challenges, many AMF species are well-adapted to such stresses and play a vital role in helping plants access nutrients in harsh environments. Therefore, these soil characteristics likely influence AMF species and how effectively they dominate. The results which are presented in Table 2, Table 3, Table 4 and Table 5 highlight the interplay between rhizosphere dynamics, soil chemical characteristics, and the distribution and abundance of arbuscular mycorrhizal fungi (AMF) across different plant species during the summer and winter seasons. The analysis underscores the critical role of the rhizosphere environment in shaping AMF colonization and community diversity, demonstrating how seasonal variations and plant-specific interactions influence fungal presence and activity.

Table 1. Soil physicochemical characteristics.

Table 2. Soil chemical characteristics of rhizosphere plants during the summer season.

Table 3. Soil chemical characteristics of rhizosphere plants during the winter season.

Table 4. Spore count and AMF colonization percentage of different plants during summer season.

Table 5. Spore count and AMF colonization percentage of different plants during winter season.

3.1. Rhizosphere Soil Nutrients

Soil nutrient composition, as shown in Table 2 and Table 3, varied significantly among plant species and across seasons, reflecting species-specific rhizosphere dynamics. Nitrogen (N) levels, in particular, differ significantly among plant species and seasons, showing the impact of plant-specific rhizosphere dynamics. Nitrogen (N) levels were the highest in Phragmites australis during summer (437.8 mg/kg) and winter (482.5 mg/kg), likely due to its dense root system enhancing microbial activity and nutrient cycling. Conversely, Calotropis procera showed the lowest nitrogen levels (70.7 mg/kg in summer and 72 mg/kg in winter), possibly due to its adaptation to nutrient-poor soils and the inhibitory effects of its allelopathic activity on microbial and AMF colonization. These results (as given in Table 2 and Table 3) show a close link between plant traits, rhizosphere processes, and soil nutrients.

Phosphorus (P) availability was generally low in summer, ranging from 1.3 mg/kg in Phragmites australis to 0.1 mg/kg in Calotropis procera. In winter, P levels ranged from 1.5 mg/kg to 0.1 mg/kg for the same species (Table 2 and Table 3). On the other hand, the study revealed that potassium (K), sodium (Na), calcium (Ca), and magnesium (Mg) concentrations showed notable variability, reflecting differences in soil type, plant exudates, and microbial interactions, particularly, arbuscular mycorrhizal fungi (AMF) during the summer and winter seasons (Table 2 and Table 3). Both total organic carbon (TOC) and organic matter (OM) showed seasonal variations in Phragmites australis, with lower values recorded during summer (2.2% for TOC and 3.6% for OM). These values increased slightly in winter, reaching 2.1% for TOC and 3.4% for OM, possibly due to enhanced microbial activity under optimal soil temperature conditions. In contrast, Calotropis procera consistently exhibited lower TOC and OM values, with summer measurements of 0.53% and 0.5%, respectively, and a slight increase in winter to 0.6% for TOC and 1.41% for OM. This seasonal variation highlights the influence of temperature and microbial activity on soil organic content.

3.2. AMF Spore Count and Community Diversity

The results, as shown in Table 4 and Table 5, highlight the influence of seasonal variation and rhizosphere dynamics on AMF spore counts. In summer, Phragmites australis and Cyperus rotundus recorded the highest spore counts, with 175 and 162 spores/100 g of soil, respectively. In winter, their counts decreased to 124 and 116 spores/100 g. In contrast, Calotropis procera consistently showed the lowest spore counts in both seasons—101 spores/100 g in summer and 63 spores/100 g in winter.

3.3. AMF Spore Diversity

Shannon biodiversity indices of AMF spores revealed that the values ranged between 0.692 for Zygophyllum simplex and 0.653 for Salsola imbricata, whereas Simpson biodiversity indices of AMF spores recorded values that ranged between 0.498 for Zygophyllum simplex and Citrullus colocynthis and 0.461 for Salsola imbricate (Table 6). Overall, the findings concluded that a wide range of AMF spore species was identified and isolated from various wild plant rhizosphere zones, including Glomus ambisporum, Claroideoglomus etunicatum, Rhizophagus intraradices, and Diversispora globifera; in the same context, it is notable that the G. ambisporum was the most common among different plant rhizospheres which may reflect how rhizosphere dynamics and root attraction influence the arbuscular mycorrhizal fungi (AMF) community, which varied among the studied plant species, indicating differences in symbiotic associations (Table 6 and Figure 1).

Table 6. Shannon–Simpson biodiversity index of AMF spores isolated from the rhizosphere of different plant species during the summer and winter seasons.

Figure 1. AMF spores isolated from a various wild plants during summer and winter: (a) intact spore of Glomus ambisporum, (b) crashed spore of Rhizophagus intraradices, (c) crashed spore of Claroideoglomus etunicatum, (d) crashed spore of Rhizophagus fasciculatus, (e) crashed spore of Diversispora globifera, (f) crashed spore of Funneliformis geosporume, (g) crashed spore of Funneliformis mosseae, (h) crashed spore of Gigaspora spp. Scale bar: 100 µm.

3.4. AMF Structure Colonization Percentage % (Mycelium, Vesicles, and Arbuscules)

Regarding Mycelium percentage, the results revealed that the Phragmites australis recorded a higher value of Mycelium percentage at 90% compared to 57% for Calotropis procera in summer; the Mycelium percentage recorded a lower value at 84% and 55% for Phragmites australis and Calotropis procera, respectively, in winter (Table 4 and Table 5).

3.5. Vesicles and Arbuscules

The findings showed that the percentage of vesicles in Phragmites australis roots increased noticeably to 36% in summer, followed by a significant decline to 24% in winter. In contrast, Calotropis procera exhibited a consistently lower vesicle percentage, declining significantly to 22% in summer and increasing to 42% in winter. Additionally, ANOVA analysis revealed that the percentage of arbuscules in Phragmites australis increased significantly to 76% in summer, but declined to 69% in winter (Table 7). On the other hand, Calotropis procera showed a significant decrease in arbuscule percentage to 49%, followed by a noticeable increase to 52% in winter (Table 4 and Table 5). The colonization structures of arbuscular mycorrhizal fungi (AMF), including hyphae (H), arbuscules (A), vesicles (V), and spores (S), were observed in the roots of various wild plant species, as shown in Figure 2. In Phragmites australis, spores, hyphae, and vesicles were found. Citrullus colocynthis had hyphae and vesicles. Malva parviflora showed hyphae and arbuscules. Calotropis procera had spores, hyphae, and vesicles. Pelargonium peltatum had hyphae and spores, and Cyperus rotundus had hyphae and vesicles. In Solanum nigrum, spores, hyphae, and vesicles were present, while Zygophyllum simplex showed spores and hyphae.

Table 7. Pearson correlation between soil properties and AMF spores during winter and summer seasons.

Figure 2. Colonization structures of arbuscular mycorrhizal fungi (AMF), including hyphae, arbuscules, vesicles, and spores, observed in roots of various wild plants. AMF spores, hyphae, and spores were present in Phragmites australis roots (1); hyphae and vesicles in Citrullus colocynthis (2); hyphae and arbuscules in Malva parviflora (3); spores, hyphae, and vesicles in Calotropis procera (4); hyphae and spores in Pelagonium peltatum (5); hyphae and vesicles in Cyperus rotundus (6); spores, hyphae, and vesicles in Solanum nigrum (7); and spores and hyphae in Zygophyllum simplex (8). Scale bar: 100 µm. Abbreviations: H—Hyphae; S—Spore; V—Vesicle; A—Arbuscular.

The results showed that the number of AMF spores was strongly correlated to several soil nutrients and organic matter. Total organic carbon (TOC) and organic matter (OM) had the highest positive correlations with spore count in both summer and winter. This means that soils with more organic material tend to have more AMF spores, likely because organic matter improves soil health and supports fungal growth. Nutrients like phosphorus and nitrogen also showed strong positive correlations, especially in the summer, which may be due to higher plant activity during this time. Sodium had an unusually high correlation in summer, which could be related to how certain plants and fungi adapt to salty soils. On the other hand, factors like pH, salinity (EC), and soil texture (sand, silt, and clay) showed little or no correlation with AMF spore count. This suggests that organic matter and key nutrients are more important than soil structure in supporting AMF abundance

Our findings show that soil properties play a major role in shaping AMF spore density and distribution in the rhizosphere. These differences directly affect the chances of root infection and colonization. This agrees with Janowski and Leski (2022), who pointed out that fungal distribution in soils is largely controlled by soil chemistry, plant interactions, and dispersal pathways. We also found that nutrient availability depends not only on soil conditions but also on root system density and the presence of AMF spores and colonization. Similar observations were reported by Adil et al. (2017), who showed that both seasonal changes and host traits influence colonization intensity and spore abundance [,].

The Poaceae (Gramineae) family stood out in our study for its strong ability to form mutualistic associations with AMF. This is likely due to their dense, fibrous root systems, which create ideal conditions for fungal growth. Our results are consistent with Ahulu et al. (2005), who found that most Poaceae species are naturally mycorrhizal. In contrast, Calotropis procera consistently showed the lowest values for colonization and spore counts. This may be explained by its allelopathic properties and aggressive growth behavior, which can suppress AMF activity [].

Seasonal variation was another clear driver of AMF dynamics. Colonization rates and spore counts were lower in winter but rose sharply in summer. Warmer conditions likely provide more favorable soil temperatures and physicochemical traits for fungal activity. These results align with Tshwene-Mauchaza and Aguirre-Gutiérrez (2019), who highlighted the importance of climate—especially temperature and rainfall—in shaping AMF distribution and seasonal tolerance []. In addition, the results showed that in summer, almost all nutrients (N, P, K, Ca, Mg, Na) were strongly linked to spore counts. Sodium and total organic carbon (TOC) had the highest correlations, showing that when plants and microbes are most active, nutrients and carbon directly support spore production. Organic matter (OM) also showed a very strong relationship with spores. However, in winter, the pattern was different. TOC and OM still had strong correlations with spores, but most nutrients (especially K, Ca, and Mg) were less important. Nitrogen and phosphorus still influenced spores, but not as much as in summer. This suggests that in winter, when plant activity is lower, AMF rely more on organic matter and carbon in the soil rather than fresh nutrient availability. Overall, organic matter is important for AMF in both seasons, but nutrients play a bigger role during summer when conditions are better for growth.

Finally, our study suggests that declines in AMF colonization may also be linked to root architecture and nutrient levels at deeper soil layers. This pattern has been widely reported in earlier research (Jakobsen and Nielsen, 1983; Smith, 1978; Sutton, 1973; Sutton and Barron, 1972), which showed that both spore density and hyphal infection decrease with soil depth. Similar declines in colonized root length and spore counts were documented by Koide and Mooney (1987) and Zajicek et al. (1986). More recent studies (Higo et al., 2013; Oehl et al., 2005; Muleta et al., 2008; Säle et al., 2015) also support this trend, confirming that AMF abundance, spore density, and root colonization are generally reduced in deeper soil layers [,,,,,,,,,].

4. Conclusions

The study highlights the complex relationships between soil properties, seasonal changes, plant type, and the abundance and activity of arbuscular mycorrhizal fungi (AMF) in arid environments. Despite challenging soil physicochemical properties such as sandy loam texture, alkalinity, moderate salinity, and high CaCO₃ content, the AMF were able to establish, sporulate, and form symbiotic associations with a range of local wild plants. Arid conditions usually limit nutrient availability and fungal growth, but they did not completely stop AMF activity. This shows how tough and adaptable these fungi are in harsh environments. One important finding of this study is that organic matter and soil nutrients play a major role in affecting the number of AMF spores. Both total organic carbon (TOC) and organic matter (OM) exhibited strong positive correlations with spore counts across seasons, underlining the importance of soil organic content in maintaining and enhancing fungal populations.

Nutrients like phosphorus and nitrogen, especially in summer, helped increase AMF activity. This is likely because plants grow more and have higher metabolism, which helps fungi develop. These results show that organic matter and nutrients matter more than soil texture or salinity for AMF growth in dry soils. Soil pH, electrical conductivity (EC), and texture (sand, silt, and clay) are usually important for soil microbes, but they showed little or no effect on AMF spore density here. This may imply that AMF communities in arid environments have adapted to a broad range of physical soil characteristics and are more sensitive to biological and chemical properties, especially organic inputs and nutrient cycling. Phragmites australis and Cyperus rotundus consistently supported higher nutrient levels, spore densities, and root colonization rates, positioning them as favorable partners for AMF in degraded or nutrient-deficient environments. In contrast, Calotropis procera showed the lowest AMF colonization, nutrient levels, and spore counts, potentially due to its allelopathic properties or association with less favorable soil conditions. These plant-specific differences highlight the importance of host selection in ecological restoration and sustainable land management, particularly in arid and semi-arid regions where soil fertility is a major constraint.

Seasonal variability also played a significant role in AMF dynamics, with summer months exhibiting greater AMF activity compared to winter. This is likely linked to enhanced plant growth and increased soil microbial activity under warmer conditions, which provide better opportunities for AMF sporulation and colonization. The dominance of widely distributed AMF species such as Glomus ambisporum, Rhizophagus intraradices, and Claroideoglomus etunicatum suggests that a core group of AMF taxa are capable of thriving in a range of plant–soil environments, albeit with varying levels of efficiency depending on host and season. Future research should clarify the roles of different AMF species in arid ecosystems under changing climates and diverse plants. Long-term studies can show how AMF respond to drought and heat. Molecular tools can reveal hidden diversity and functions. Studying AMF interactions with other microbes and testing organic amendments or bioinoculants may help improve soil health and sustainable farming in arid regions.

Author Contributions

Conceptualization, S.S.A., F.A. and S.N.S.; Formal analysis, S.S.A. and F.A.; Data curation, F.A., S.S.A. and S.N.S.; Writing—original draft preparation, B.A., S.S.A. and F.A.; Writing—review and editing, S.S.A., F.A., B.A. and S.S.A.; Supervision, S.S.A. All authors have read and agreed to the published version of the manuscript.

Funding

This project was funded by Ongoing Research Funding program, King Saud University, Riyadh, Saudi Arabia. ORF-2025-866.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The data used in this study are available from the corresponding author upon reasonable request.

Acknowledgments

The authors extend their deep appreciation to King Saud University, Riyadh, Saudi Arabia for funding the Ongoing Research Funding program, (ORF-2025-866).

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. AMF spores isolated from a various wild plants during summer and winter: (a) intact spore of Glomus ambisporum, (b) crashed spore of Rhizophagus intraradices, (c) crashed spore of Claroideoglomus etunicatum, (d) crashed spore of Rhizophagus fasciculatus, (e) crashed spore of Diversispora globifera, (f) crashed spore of Funneliformis geosporume, (g) crashed spore of Funneliformis mosseae, (h) crashed spore of Gigaspora spp. Scale bar: 100 µm.

Figure 2. Colonization structures of arbuscular mycorrhizal fungi (AMF), including hyphae, arbuscules, vesicles, and spores, observed in roots of various wild plants. AMF spores, hyphae, and spores were present in Phragmites australis roots (1); hyphae and vesicles in Citrullus colocynthis (2); hyphae and arbuscules in Malva parviflora (3); spores, hyphae, and vesicles in Calotropis procera (4); hyphae and spores in Pelagonium peltatum (5); hyphae and vesicles in Cyperus rotundus (6); spores, hyphae, and vesicles in Solanum nigrum (7); and spores and hyphae in Zygophyllum simplex (8). Scale bar: 100 µm. Abbreviations: H—Hyphae; S—Spore; V—Vesicle; A—Arbuscular.

Table 1. Soil physicochemical characteristics.

Parameter	Mean
pH (1:1)	8.1
EC dSm⁻¹	2.8
OM %	1.03
CaCO₃ %	11.91
Sand %	58
Silt %	23
Clay %	19
CEC cmol·kg⁻¹	5.81

Table 2. Soil chemical characteristics of rhizosphere plants during the summer season.

Plant Species	N	P	K	Na	Ca	Mg	Total Organic Carbon (TOC)	Organic Matter (OM)
Solanum nigrum	122 ± 2.9 e	0.9 ± 0.02 bc	5.6 ± 0.26 ef	127 ± 3 e	2.7 ± 0.2 d	0.6 ± 0.03 de	1.2 ± 0.2 cd	2.2 ± 0.12 bc
Zygophyllum coccineum	79.9 ± 0.9 f	0.6 ± 0.09 de	2.4 ± 0.09 gh	9.4 ± 0.36 fgh	1.5 ± 0.14 d	0.5 ± 0.04 de	0.79 ± 0.06 efg	1.4 ± 0.27 cd
Phragmites australis	437.8 ± 8.7 a	1.3 ± 0.1 a	92.7 ± 2.7 a	312.6 ± 7 a	54.9 ± 2.8 a	20.6 ± 0.04 a	2.1 ± 0.1 a	3.4 ± 0.35 a
Zygophyllum simplex	119.8 ± 0.9 e	0.7 ± 0.02 de	2.9 ± 0.4 fgh	16.8 ± 0.4 fg	2.5 ± 0.19 d	0.5 ± 0.02 de	1 ± 0.03 def	1.4 ± 0.02 cd
Malva parviflora	221.2 ± 1.4 c	1.2 ± 0.02 a	14.5 ± 0.35 c	251.4 ± 2.5 c	9.4 ± 0.4 c	4.4 ± 0.6 c	1.5 ± 0.1 b	2.6 ± 0.29 ab
Cyperus rotundus L.	233.6 ± 7.4 b	1.2 ± 0.05 a	67.3 ± 2 b	276.9 ± 7 b	31.4 ± 0.98 b	17 ± 0.06 b	1.7 ± 0.3 b	2.7 ± 0.22 ab
Calotropis procera	70.7 ± 1.1 f	0.1 ± 0.006 f	1.3 ± 0.1 h	0.6 ± 0.03 h	1.2 ± 0.09 d	0.2 ± 0.02 e	0.53 ± 0.04 g	0.5 ± 0.2 d
Pelargonium peltatum	208.8 ± 0.97 d	1.1 ± 0.06 ab	8.6 ± 0.28 d	164.6 ± 4.6 d	3.1 ± 0.09 d	0.9 ± 0.07 d	1.5 ± 0.1 bc	2.6 ± 0.17 ab
Salsola imbricata	74.6 ± 1.8 f	0.5 ± 0.1 de	2.2 ± 0.08 gh	5.1 ± 0.1 gh	1.3 ± 0.03 d	0.3 ± 0.02 de	0.7 ± 0.04 fg	1.3 ± 0.5 cd
Citrullus colocynthis	121.5 ± 2.8 e	0.8 ± 0.04 cd	4.8 ± 0.35 efg	19.7 ± 1.3 f	2.5 ± 0.09 d	0.6 ± 0.04 de	1.1 ± 0.1 de	1.8 ± 0.26 bc
Senna italica	208.2 ± 3.3 d	0.9 ± 0.1 c	5.8 ± 0.37 de	153.6 ± 9.9 d	2.9 ± 0.16 d	0.6 ± 0.04 de	1.2 ± 0.1 cd	2.4 ± 0.8 ab

The values with the same letters are not significantly different from each other according to the LSD test (p < 0.05).

Table 3. Soil chemical characteristics of rhizosphere plants during the winter season.

Plant Species	N	P	K	Na	Ca	Mg	Total Organic Carbon (TOC)	Organic Matter (OM)
Solanum nigrum	130.2 ± 1.2 e	1.1 ± 0.02 c	6. ± 0.4 e	139.8 ± 1.4 f	2.7 ± 0.07 d	0.6 ± 0.03 d	1.4 ± 0.1 cd	2.7 ± 0.1 ab
Zygophyllum coccineum	82.7 ± 1.2 g	0.7 ± 0.02 de	2.6 ± 0.1 fg	11.5 ± 0.5 h	1.6 ± 0.14 d	0.6 ± 0.03 d	0.9 ± 0.02 def	1.5 ± 0.7 cd
Phragmites australis	482.5 ± 4.8 a	1.5 ± 0.2 a	98. ± 2.6 a	323.8 ± 2 a	58.6 ± 3.7 a	22.2 ± 0.7 a	2.2 ± 0.03 a	3.6 ± 0.2 a
Zygophyllum simplex	123.5 ± 1.43	0.8 ± 0.1 d	3.04 ± 0.1 fg	17.6 ± 0.8 h	2.7 ± 0.02 d	0.6 ± 0.1 d	1.24 ± 0.03 cde	2.2 ± 0.1 cd
Malva parviflora	225.9 ± 0.98 c	1.3 ± 0.03 b	15.8 ± 0.17 c	259.9 ± 1 c	14.3 ± 0.24 c	5.6 ± 0.2 c	1.7 ± 0.1 abc	3.01 ± 0.8 ab
Cyperus rotundus L.	254.8 ± 1.6 b	1.3 ± 0.1 b	70.7 ± 1.5 b	285.8 ± 2.9 b	35.61 ± 0.9 b	18.6 ± 0.8 d	1.9 ± 0.1 ab	3.1 ± 0.2 ab
Calotropis procera	72 ± 1 h	0.13 ± 0.02 f	1.4 ± 0.2 g	0.7 ± 0.1 j	1.3 ± 0.03 d	0.3 ± 0.01 d	0.6 ± 0.3 f	1.41 ± 0.1 d
Pelargonium peltatum	211.1 ± 1 d	1.24 ± 0.02 bc	8.9 ± 0.3 d	174.5 ± 1.3 d	3.3 ± 0.1 d	1.1 ± 0.04 d	1.7 ± 0.2 bc	2.9 ± 0.2 ab
Salsola imbricata	78.7 ± 1 g	0.6 ± 0.02 e	2.31 ± 0.1 g	5.4 ± 0.2 i	1.3 ± 0.02 d	0.4 ± 0.01 d	0.84 ± 0.4 ef	1.48 ± 0.7 cd
Citrullus colocynthis	124.4 ± 1.1 f	0.8 ± 0.1 d	5. ± 0.2 ef	20.4 ± 0.4 g	2.7 ± 0.02 d	0.6 ± 0.04 d	1.3 ± 0.1 cde	2.6 ± 0.14 abc
Senna italica	211. ± 1 d	1.23 ± 0.1 bc	6.21 ± 0.2 de	166.3 ± 2 e	3.14 ± 0.13 d	0.7 ± 0.02 d	1.7 ± 0.03 bc	2.9 ± 0.02 ab

The values with the same letters are not significantly different from each other according to the LSD test (p < 0.05).

Table 4. Spore count and AMF colonization percentage of different plants during summer season.

Plant Species	Spore Count	Myciilum %	Vesicles %	Arbuscules %
Solanum nigrum	125.7 ± 5.7 d	82.7 ± 2.6 abc	29 ± 9.8 a	62.3 ± 4.6 abc
Zygophyllum coccineum	117.3 ± 2.3 d	75.3 ± 4.6 c	22.7 ± 4.3 a	55.6 ± 5.9 abc
Phragmites australis	175 ± 5.3 a	90 ± 1.7 a	36 ± 5.9 a	76 ± 4.7 a
Zygophyllum simplex	120 ± 4.7 d	77.3 ± 2.9 bc	24.3 ± 6.9 a	56.7 ± 11.3 abc
Malva parviflora	157.7 ± 2.9 b	86 ± 4.3 abc	34.3 ± 6.9 a	69.3 ± 4.5 abc
Cyperus rotundus L.	162 ± 3.2 ab	87.7 ± 2.9 ab	34.7 ± 4.6 a	70 ± 8.9 ab
Calotropis procera	101.3 ± 7.9 e	57.7 ± 4.7 d	22.3 ± 4.7 a	49 ± 9.8 c
Pelargonium peltatum	155.3 ± 3.8 bc	85.3 ± 4.9 abc	33.3 ± 3.8 a	65.3 ± 5.6 abc
Salsola imbricata	114.3 ± 4 de	74.3 ± 4.7 c	22.3 ± 4.7 a	52.3 ± 2.9 bc
Citrullus colocynthis	122 ± 2.9 d	82.7 ± 2.6 abc	24.7 ± 6.2 a	57 ± 5.7 abc
Senna italica	142.3 ± 4.4 c	83.3 ± 5 abc	32.3 ± 9.8 a	64.6 ± 6.2 abc

The values with the same letters are not significantly different from each other according to the LSD test (p < 0.05).

Table 5. Spore count and AMF colonization percentage of different plants during winter season.

Plants Species	Spore Count	Myciilum %	Vesicles %	Arbuscules %
Solanum nigrum	108.7 ± 13.7 abc	81 ± 2.8 a	35.7 ± 5.9 ab	62.7 ± 8 ab
Zygophyllum coccineum	67 ± 7.8 bc	75 ± 4.6 a	42.3 ± 7.9 a	55.7 ± 5.9 ab
Phragmites australis	124 ± 5 a	84.7 ± 4.4 a	24.3 ± 4.7 b	69.3 ± 4.5 a
Zygophyllum simplex	107.3 ± 3.2 abc	75.7 ± 5.9 a	37.7 ± 2.3 ab	55.7 ± 5.9 ab
Malva parviflora	113 ± 6 abc	84 ± 3 a	29.3 ± 2.3 ab	65.3 ± 6.7 ab
Cyperus rotundus L.	116.7 ± 10.8 ab	84.3 ± 5.9 a	24.3 ± 5.9 b	65.7 ± 5.5 ab
Calotropis procera	63 ± 2.8 c	55 ± 1.2 b	42.7 ± 4.3 a	52.7 ± 8.7 b
Pelargonium peltatum	111.7 ± 8.4 abc	82. ± 8.7 a	31.3 ± 4.3 ab	64.7 ± 5.6 ab
Salsola imbricata	64. ± 4 c	74 ± 5.8 a	42.7 ± 2.9 a	54.3 ± 4.6 ab
Citrullus colocynthis	107.7 ± 5.2 abc	75.7 ± 5.9 a	35.7 ± 5.9 ab	59 ± 6.1 ab
Senna italica	109.7 ± 4 abc	82 ± 3.2 a	34.7 ± 6.9 ab	64.7 ± 8 ab

The values with the same letters are not significantly different from each other according to the LSD test (p < 0.05).

Table 6. Shannon–Simpson biodiversity index of AMF spores isolated from the rhizosphere of different plant species during the summer and winter seasons.

Plant Species	Spore Count/100 gm Dry Soil		Shannon (H’)	Simpson (D)	AMF Spore Diversity
Plant Species	Summer	Winter	Shannon (H’)	Simpson (D)	AMF Spore Diversity
Solanum nigrum	125	108	0.690	0.497	Glomus ambisporum, Rhizophagus intraradices, Claroideoglomus etunicatum
Zygophyllum coccineum	117	67	0.656	0.463	Diversispora globifera, Funneliformis geosporum, Rhizophagus intraradices, Claroideoglomus etunicatum, Glomus ambisporum
Phragmites australis	175	124	0.679	0.485	Rhizophagus intraradices, Funneliformis mosseae, Glomus ambisporum, Claroideoglomus etunicatum, Diversispora globifera, Gigaspora spp.
Zygophyllum simplex	120	107	0.692	0.498	Diversispora globifera, Funneliformis mosseae, Rhizophagus fasciculatus, Glomus ambisporum, Claroideoglomus etunicatum
Malva parviflora	157	113	0.680	0.487	Funneliformis mosseae, Glomus ambisporum, Diversispora globifera, Claroideoglomus etunicatum, Rhizophagus fasciculatus
Cyperus rotundus L.	162	116	0.679	0.486	Diversispora globifera, Funneliformis geosporum, Glomus ambisporum, Rhizophagus intraradices, Claroideoglomus etunicatum
Calotropis procera	101	63	0.666	0.473	Rhizophagus intraradices, Diversispora globifera, Glomus ambisporum, Claroideoglomus etunicatum, Rhizophagus fasciculatus
Pelargonium peltatum	155	111	0.679	0.486	Claroideoglomus etunicatum, Diversispora globifera
Salsola imbricata	114	64	0.653	0.461	Gigaspora spp., Glomus ambisporum, Claroideoglomus etunicatum
Citrullus colocynthis	122	107	0.691	0.498	Rhizophagus intraradices, Claroideoglomus etunicatum, Diversispora globifera, Glomus ambisporum
Senna italica	142	109	0.684	0.491	Funneliformis mosseae, Claroideoglomus etunicatum

Table 7. Pearson correlation between soil properties and AMF spores during winter and summer seasons.

Variables	Summer Seasons	Winter Seasons
Variables	AMF Spore Count
N (mg·kg⁻¹)	0.91 **	0.71 **
P (mg·kg⁻¹)	0.94 **	0.87 **
K (mg·kg⁻¹)	0.76 **	0.53 *
Na (mg·kg⁻¹)	0.96 **	0.76 **
Ca (mg·kg⁻¹)	0.75 **	0.53 *
Mg (mg·kg⁻¹)	0.76 **	0.52 *
Total Organic Carbon (TOC) (mg·kg⁻¹)	0.96 **	0.92 **
Organic Matter (OM) %	0.96 **	0.96 **
pH	0.05	0.02
Sand %	−0.31	0.16
Silt %	0.17	0.07
Caly %	−0.05	−0.36
EC (dSm⁻¹)	−0.03	−0.39
CEC (cmol·kg⁻¹)	0.41	0.32
CaCO₃ %	0.07	−0.11
Spore Count	1.00	1.00

* Correlation is significant at p = 0.05. ** Correlation is significant at p = 0.01.

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