Seasonal Dynamics of Physical, Hydraulic, and Physico-Chemical Attributes of the Soil across Altitudinal Gradients in the Alpine Wetlands of Lesotho

Seasonal Dynamics of Physical, Hydraulic, and Physico-Chemical Attributes of the Soil across Altitudinal Gradients in the Alpine Wetlands of Lesotho | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Seasonal Dynamics of Physical, Hydraulic, and Physico-Chemical Attributes of the Soil across Altitudinal Gradients in the Alpine Wetlands of Lesotho Knight Nthebere, Mosiuoa Mochala This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9212062/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Seasonal fluctuations significantly influenced soil physical, hydraulic, and physico-chemical properties in Lesotho’s alpine wetlands, yet these ecosystems remain largely understudied. This study evaluated composite seasonal index and examined soil variation and seasonal changes in soil attributes across altitudes (2500–3155 m a.s.l.), equivalent to wetlands from three sub-catchments (blocks): Khorong and Tenesolo (Senqunyane), Khamoqana and Khalong-la-Lichelete (Sani), and Lets’eng-la-Likhama and Koting-Sa-ha Ramosetsana (Khubelu), during the standard (spring, September 2024) and peak (summer, February 2025) wet seasons. The soil samples were collected in September 2024 (standard) and February 2025 (peak) wet seasons and analyzed for bulk density (BD), saturated hydraulic conductivity (Ksat), infiltration rate (IR), water holding capacity (WHC), texture, pH, electrical conductivity (EC), cation exchange capacity (CEC) and soil organic carbon (SOC) following standard procedures. Soil texture exhibited clear altitudinal trends: sand decreased from 64.97% at lower elevations to 39.79% upslope, whereas silt and clay increased, resulting in sandy-loam at lower and loam at higher sites. Seasonal variations, though subtle, were measurable: sand (± 0.2%), silt (± 1%), clay (± 1.3%), BD (6.8%), Ksat (13%), IR (5.1%), and WHC (2.6%). SOC rose slightly (~ 0.4%) in summer, peaking at Koting-Sa-ha Ramosetsana. Correlation analysis indicated strong negative relationships between sand and both clay and silt, while BD inversely related to IR. Positive associations were observed between WHC and Ksat, and IR correlated closely with SOC. Overall, these results underscore that seasonal hydrological dynamics and elevation jointly shape soil structure, water movement, and carbon storage in Lesotho’s alpine-wetlands, emphasizing their ecological sensitivity and need for targeted conservation. Agroecology Seasonal soil dynamics seasonal soil index soil attributes climate adaptation ecosystem-based adaptability high altitudinal gradient Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Alpine wetlands are increasingly acknowledged as ecologically fragile yet functionally indispensable components of mountain environments, where they modulate hydrological fluxes, conserve biodiversity, and attenuate climate variability (IPCC, 2021; Mitsch et al., 2015 ). Within the high-elevation landscapes of Lesotho, these wetlands constitute critical headwater systems that regulate stream discharge, facilitate groundwater recharge, and safeguard downstream water supplies, including key transboundary water resources. Emerging evidence indicates that, despite occupying relatively small spatial extents, mountain wetlands exert disproportionately large influences on watershed hydrology and carbon sequestration (Lin et al., 2023 ; Nthebere et al., 2025 ). The ecological performance of alpine wetlands is fundamentally regulated by soil physical, hydraulic, and physico-chemical characteristics, which collectively influence water retention, infiltration dynamics, aeration, nutrient turnover, and carbon stabilization. Consequently, examining the temporal variability of these soil attributes, particularly across seasons, is essential for advancing sustainable wetland management and enhancing climate resilience in highland systems (Nthebere et al., 2025 ). Soil physical properties, such as bulk density, total porosity, texture, and aggregate stability, determine structural coherence, resistance to erosion, and root penetration. Hydraulic attributes, including infiltration rate, saturated hydraulic conductivity, and maximum water holding capacity, regulate soil-water interactions, influencing runoff generation and aquifer recharge. Key physico-chemical indicators, notably soil pH, electrical conductivity and soil organic carbon (SOC), govern microbial activity and biogeochemical cycling that sustain ecosystem productivity (Lal, 2020 ; Lehmann et al., 2020 ). These interrelated properties are dynamic, varying with seasons, vegetation across altitude (Mirza and Patil, 2020 ). Elevation-driven changes in plant communities significantly influence SOC stabilization and soil hydraulic behavior (Zhang et al ., 2022). Soil characteristics are further shaped by interactions among abiotic factors; temperature, precipitation, topography and biotic components, including plants and microbes, which fluctuate spatially and seasonally (Das et al, 2025 ; Liu et al., 2023 ). Seasonal transitions, particularly from the standard wet season to the peak wet season, modify soil moisture, redox status, hydraulic conductivity, and nutrient availability. Progressive wetting may enhance aggregate stability and microbial mineralization, whereas sustained saturation can reduce aeration, destabilize soil structure, limit infiltration, and promote denitrification and methane production (Lin et al., 2023 ). Altitudinal gradients mediate these effects by influencing thermal regimes, rainfall intensity, vegetation composition, and decomposition dynamics, yet these interactions remain poorly documented in southern African alpine wetlands. Despite their ecological and hydrological importance, Lesotho’s alpine wetlands are rarely studied with integrated assessments that combine seasonal variability, altitudinal differentiation, and soil physical, hydraulic, and physico-chemical indicators. Most studies focus on single properties or seasons, overlooking interactive effects critical under increasing climate variability and anthropogenic pressures such as overgrazing and land-use intensification (IPCC, 2021). This study addresses the central question: How do seasonal transitions from the standard to peak wet season, across altitudinal gradients, influence the integrated soil functionality of Lesotho’s alpine wetlands? It is hypothesized that seasonal progression significantly alters soil physical, hydraulic, and physico-chemical properties, with systematic variation along elevation gradients. The study aimed to quantify seasonal changes in soil attributes across elevation zones during both wet-season phases and to evaluate a composite seasonal index of selected soil properties reflecting functional shifts along altitudinal gradients. Assessing soil variability via an integrated index is vital because alpine wetlands regulate watershed hydrology, influence SOC stabilization and greenhouse gas fluxes, and provide a quantitative framework to detect changes that individual indicators may miss. Integrating seasonal and altitudinal controls enhances predictive capacity under climate change and supports sustainable wetland management in Lesotho’s mountainous landscapes. By combining seasonal dynamics with elevation-driven controls through a holistic soil quality framework, this study offers a comprehensive approach to understanding soil functionality and ecological resilience in alpine wetlands, informing climate adaptation and watershed management strategies. Materials and Methods 2.1. Description of the Study Area The investigation was conducted in the high-altitude regions of Lesotho, spanning elevations between 2500 and 3089 m above sea level within the mountain agro-ecological zones (AEZs) of Mokhotlong and Ha-Mohale. The study focused on three sub-catchments; Khubelu, Senqunyane, and Sani, situated within the Upper Senqu main catchment. Two alpine wetlands were identified and sampled in each sub-catchment, giving a total of six study wetlands. In the Khubelu sub-catchment, the Lets’eng-la-Likhama and Koting-sa-ha Ramosetsana wetlands were included. The Senqunyane sub-catchment comprised the Khorong and Tenesolo wetlands, while the Sani sub-catchment encompassed the Khamoqana and Khalong-la-Lichelete wetlands. Figure 1 presents a map of Lesotho, indicating its ten administrative districts within the African continent and highlighting the selected alpine wetland sites. Figure 2 illustrates the GPS-based layout of the Upper Senqu main catchment, showing the location of the three sub-catchments; Khubelu, Senqunyane, and Sani, and the distribution of the sampled wetlands within them. 2.2 The Alpine Wetland Features Across the alpine wetlands, human settlement is sparse, with activity largely confined to seasonal cattle posts. Within Lesotho’s highland agro-ecological zones, vegetation is predominantly composed of shrubs and grasses adapted to high-altitude conditions. The underlying geology of these wetlands is mainly associated with formations described as Lesotho Genesis (Schmitz and Rooyani, 1987 ). At Tenesolo, the wetland showed pronounced degradation, largely attributable to repeated vegetation burning, past overgrazing, and extensive burrowing by ice rats (Supplementary Figure S1a). In contrast, land use around Khorong was primarily characterized by cultivation (Supplementary Figure S1b). In Khamoqana, burrowing by ice rats resulted in numerous surface holes, accelerated soil loss through runoff, and the dominance of sparse vegetation with shallow root systems (Supplementary Figure S1c). Khalong-La-Lichelete was distinguished by the occurrence of deep-rooted shrub species (Supplementary Figure S1d). The Lets’eng-La-Likhama wetland was mainly affected by limited grass cover, erosion linked to inadequate vegetative protection, and heavy grazing pressure from sheep (Supplementary Figure S1e). At Koting-Saha Ramosetsana, shrubs predominated in the surrounding landscape, while grasses were more common within the wetland itself (Supplementary Figure S1f). Soil degradation across all sites was assessed using WET-Health version 2.0, an advanced assessment framework developed to evaluate the current ecological status of wetland systems, following the approaches outlined by Kleynhans ( 1996 ) and Macfarlane et al. ( 2009 ). The assessment outcomes are presented in Table 1 , with additional detail provided in Supplementary Table S1. Table 1 Alpine wetland characteristics. Alpine Wetlands Latitude S Longitude E Soil Degradation Level Assessed with WET Health According to Kleynhans ( 1996 ) and Macfarlane et al. ( 2009 ). PES Score (%) Description Khorong −29.457168 28.268082 80 Largely natural with few modifications. A slight change in ecosystem processes is discernible, and a small loss of natural habitats and biota may have taken place. Tenesolo −29.449256 28.149214 45 Extensively altered and alterations in ecological functions accompanied by the disappearance of natural habitats and native species. Khamoqana −29.457178 28.268094 30 Seriously modified, the change in ecosystem processes, great loss of natural habitat and biota but some remaining natural habitat features are still being recognized. Khalong-La-Lichelete −29.563552 29.247207 90 Unmodified natural wetland Lets’eng-La-Likhama −29.076355 28.836095 40 Largely modified with a large change in ecosystem processes and loss of natural habitat, and biota has occurred. Koting-Sa-ha Ramosetsana −29.022686 28.871324 85 Largely natural with few modifications and a slight change in ecosystem processes being discernible and a small loss of natural habitats and biota may have taken place. S = Southern Hemisphere; E = Eastern Hemisphere; PES = Percentage. 2.3. Climate Lesotho’s climatic conditions are largely influenced by its position on the central southern African Plateau. The country experiences a sub-humid, cool temperate climate, with distinct seasonal contrasts marked by hot, wet summers and cold, dry winters (Wu et al., 2022 ). June is generally the coldest month, when mean minimum temperatures are close to 0°C. In the lowland zones, winter temperatures commonly range from about − 1°C to − 3°C, while the high-altitude regions may experience more severe conditions, with temperatures declining to between roughly − 6°C and − 8.5°C (Malebajoa, 2010 ). Average annual temperatures are approximately 15.2°C in the lowlands and about 7°C in the highlands. January typically registers the highest average maximum temperatures, reaching nearly 32°C in the lowlands and around 20°C in the highlands. Rainfall distribution varies geographically, generally ranging from about 500 mm to 1200 mm per year, with the northern and eastern parts of the country receiving comparatively higher amounts. The majority of precipitation, around 85%, occurs during the summer months from October to April. Frost and occasional snowfall are common features of winter in the mountainous areas. Mean annual rainfall in the Tenesolo and Khorong wetlands is close to 1000 mm, while slightly higher averages of approximately 1044 mm are reported for the Khamoqana, Khalong-La-Lichelete, Lets’eng-La-Likhama, and Koting-Sa-ha Ramosetsana wetlands (Malebajoa, 2010 ). 2.4. Research Design A block experimental layout was adopted, whereby differences in altitude among alpine wetlands were organized into blocks defined by catchments sharing similar attributes, particularly wetland condition (degraded versus intact) and elevation span, in order to reduce extraneous variation. Controlling for these site characteristics allowed clearer evaluation of the treatment effect, represented by altitudinal differences among wetlands. Altitude was therefore considered the main experimental factor and was represented by six alpine wetlands distributed across three sub-catchments; Khubelu, Senqunyane, and Sani, with two wetlands selected from each catchment. Specific treatment descriptions, corresponding to the altitudinal classes of the selected wetlands, are presented in Table 2 . 2.5. Wetland Selection Criteria Wetland sites were selected based on specific elevation thresholds within each catchment: ≥3000 m in Khubelu, ≥ 2500 m in Senqunyane, and ≥ 2800 m in Sani. For every catchment, two alpine wetlands were purposively selected to reflect contrasting ecological states, one exhibiting signs of degradation and the other remaining in a comparatively sound condition. The health status of the wetlands was evaluated using the WET-Health assessment tool (version 2.0) (Kleynhans, 1996 ; Macfarlane et al. 2009 ). 2.6. Selection of Sampling Points Sampling locations within each wetland were determined using a stratified random approach to ensure adequate representation of site variability. Within each stratum, a grid-based layout was applied to systematically position sampling points in alignment with the soil sampling objectives (Paulsen et al., 1991 ). To allow replication, every wetland was divided into four equal grid sections, yielding four replicate samples per site. Table 2 Treatment details. Treatment(s) Alpine Wetlands Altitude (m) asl Khorong 2500–2550 Tenesolo 2552–2600 Khamoqana 2839–2880 Khalong-La-Lichelete 2891–2995 Lets’eng-La-Likhama 3040–3080 Koting-Sa-ha Ramosetsana 3087–3155 2.7. Soil Sampling and Standard Analytical Methods Soil samples were collected at the end of September 2024 (late spring), representing the standard wet season, and again in February 2025 (summer), corresponding to the peak wet season. These two sampling periods were deliberately selected to capture intra-seasonal variability and to facilitate the development of a seasonal index reflecting shifts in soil physical, hydraulic, and physico-chemical properties under varying moisture regimes in alpine wetlands. Sampling during both phases of the wet season enabled comparison between early-season conditions, when soils begin to re-wet, and peak-season conditions, when moisture availability and hydrological activity are typically at their maximum. At each wetland, soil was collected from the 0–15 cm depth, representing the active surface layer most responsive to seasonal changes and biogeochemical processes. Within each replication, ten subsamples were taken from multiple points and thoroughly mixed to form a single composite sample. Four replications were obtained per site. The composite samples were air-dried under shade, lightly crushed, and passed through 2.0 mm and 0.5 mm sieves to obtain the required fractions for analysis. After labeling, the processed samples were stored in polyethylene bags pending laboratory evaluation. Physical, hydraulic, and physico-chemical analyses were subsequently conducted using established standard procedures (Table 3 ). Table 3 Methodology and terms of the references adopted for the analysis of physical, hydraulic and physico-chemical properties of the soil at the 0–15 cm depth. S.No Soil Property Method Reference 1 Mechanical separates Hydrometer method Gee and Or (2002) Sand (%) Silt (%) Clay (%) 2 Soil reaction (pH) Soil: water suspension (1:2.5) Jackson ( 1973 ) 3 Electrical conductivity (dS m − 1 ) 4 Bulk density (Mg m − 3 ) Core sampler Blake and Hartge ( 1986 ) 5 Soil organic carbon (g kg − 1 ) Wet oxidation Walkley and Black ( 1934 ) 6 Cation exchange capacity (cmol) (P + ) kg − 1 Sodium acetate method Bower et al. ( 1952 ) 7 Saturated hydraulic conductivity (cm hr − 1 ) Constant head method Klute and Driksen (1986) 8 Infiltration rate (cm hr − 1 ) Double-ring infiltrometer Bouwer ( 1986 ) 9 Maximum water holding capacity (%) Keen’s cup method Keen-Raczkowski (1921) Statistical Analysis The collected data were subjected to statistical evaluation using analysis of variance (ANOVA) based on a one–factor randomized block design, following the procedure outlined by Panse and Sukhatme ( 1978 ). Differences among treatment means were tested for statistical significance at the 5% probability level. Relationships among the measured soil attributes were further examined using Pearson’s correlation analysis and principal component analysis (PCA). These analyses were conducted using the SQI CAL tool (Mohanty, 2020), which is specifically developed for soil quality assessment. The analyses were performed independently for each season in order to evaluate seasonal differences in the relationships among soil parameters. Several techniques are available for determining seasonal indices; however, the simple average method was adopted in this study as described by Mirza and Patil ( 2020 ). Seasonal indices were computed to quantify temporal variability in the measured soil properties. In this context, the seasonal index (SI) for spring (representing the standard wet season) and summer (representing the peak wet season) was calculated to express the average percentage deviation of soil parameters during each season relative to the overall mean. 3.1. Method of simple averages A simple averaging approach was applied to assess seasonal variations over the time series. The seasonal index (S.I) was computed by expressing the mean value for each period as a percentage of the overall mean x i.e ., seasonal index for different periods (Mirza and Patil, 2020 ); S.I = Average of a season / Grand Average of the all seasonal averages ×100 Seasonal index for i th season, Si = Average of i th season / Grand average of k seasons × 100 Therefore, Si = x 1 / x × 100; i = 1, 2, 3… k Thus seasonal indices are, x 1 /x × 100, x 2 / x × 100,….., x k/ x × 100. Results 4.1. Seasonal variation of soil particle size distribution and texture across altitudinal gradients Soil particle size distribution (PSD) differed significantly between the standard wet and peak wet seasons along the altitudinal gradients of the selected alpine wetlands (Table 4 ). Across both seasons, sand, silt and clay contents ranged from 39.79–64.97%, 21.92–35.72% and 10.45–25.29%, respectively. Sand content showed a clear decline with increasing altitude, with significantly higher proportions recorded at Khorong, Tenesolo and Khalong-La-Lichelete, while lower values occurred at Lets'eng-La-Likhama and Koting-sa-ha Ramosetsana. In contrast, silt and clay fractions generally increased from Tenesolo to Koting-sa-ha Ramosetsana along the elevation gradient. Seasonal comparison indicated a slight increase in sand content from the standard wet season (late spring) to the peak wet season (summer), whereas clay content declined during the same period, while silt remained relatively stable. Consequently, soils at Khorong, Tenesolo and Khalong-La-Lichelete were classified as sandy loam, whereas those at Khamoqana, Lets'eng-La-Likhama and Koting-sa-ha Ramosetsana were predominantly loam in both the seasons (Table 4 ). Table 4 Seasonal variation of soil particle size distribution and texture as influenced by altitudinal gradients Treatment(s) Spring (Standard wet season) Summer (Peak wet season) Textural Class Sand Silt Clay Sand Silt Clay Wetlands Altitude (m) asl (%) Khorong 2500–2550 64.82 23.67 11.52 64.97 23.67 11.36 Sandy loam Tenesolo 2552–2600 60.48 28.92 10.61 60.63 28.92 10.45 Sandy loam Khamoqana 2839–2880 52.69 34.03 13.28 52.82 34.04 13.14 Loam Khalong-La- Lichelete 2891–2995 64.27 21.92 13.81 64.43 21.92 13.65 Sandy loam Lets'eng- La-Likhama 3040–3080 46.26 35.71 18.02 46.38 35.72 17.90 Loam Koting-Sa-ha Ramosetsana 3087–3155 39.69 35.01 25.29 39.79 35.02 25.19 Loam SE (m)± 0.079 0.004 0.082 0.279 0.140 0.245 CD (P < 0.05) 0.246 0.013 0.257 0.868 0.307 0.764 CD (P < 0.05) = Critical Difference at less than 5% probability level; SE(m) = Standard Error of the mean 4.2. Seasonal variation of soil physical and hydraulic properties across altitudinal gradients The soil physical property (bulk density) and hydraulic attributes; maximum water holding capacity, saturated hydraulic conductivity, and infiltration rate, showed significant seasonal variation along the altitudinal gradients (Table 5 ). Across the wetlands, bulk density (BD), maximum water holding capacity (MWHC), saturated hydraulic conductivity (Ksat), and infiltration rate (IR) ranged from 1.09–1.74 Mg m⁻³, 41.70–57.51%, 1.28–2.92 cm hr⁻¹, and 1.20–2.36 cm hr⁻¹, respectively. Bulk density declined with increasing altitude and was significantly lower at Koting-Sa-ha Ramosetsana (KSHM), recording 1.25 Mg m⁻³ during the standard wet season and 1.09 Mg m⁻³ during the peak wet season. In both seasons, MWHC, Ksat, and IR were significantly higher at KSHM, whereas the lowest values (p < 0.05) were observed at Tenesolo compared with the other wetlands along the altitudinal gradient (Table 5 ). Seasonal trends indicated that BD, Ksat, and IR were higher during the standard wet season but declined during the peak wet season, while MWHC increased from the standard wet season to the peak wet season across all wetlands. 4.3. Seasonal variation of soil physico-chemical properties across altitudinal gradients Soil pH was significantly influenced by altitudinal gradients across the alpine wetlands, ranging from 5.53– 6.22 in both seasons (Table 6 ). The lowest and highest pH values were recorded at Lets’eng-la-Likhama (LLL) and Khorong, respectively, indicating slightly acidic soil conditions across all wetlands. In contrast, soil electrical conductivity (EC) and cation exchange capacity (CEC) were not significantly affected by altitude, although their values varied from 0.29– 0.37 dS m⁻¹ and 20.86– 31.04 cmol (P⁺) kg⁻¹ across wetlands in both seasons. Soil organic carbon (SOC) at the 0–15 cm depth showed significant variation along the altitudinal gradient, ranging from 68.59– 95.80 g kg⁻¹. The highest SOC levels were recorded at Koting-Sa-ha Ramosetsana (KSHM) during both the standard wet season (95.03 g kg⁻¹) and the peak wet season (95.80 g kg⁻¹), while Tenesolo (TNL) exhibited the lowest SOC among the wetlands. Overall, SOC tended to increase with elevation in both seasons. Seasonally, EC, CEC, and pH showed a slight decline from the standard wet season to the peak wet season, whereas SOC increased (Table 6 ). Table 5 Seasonal variation of soil physical and hydraulic properties as influenced by altitudinal gradients Treatment(s) Spring (Standard wet season) Summer (Peak wet season) BD (Mg m − 3 ) Ksat IR MWHC (%) BD (Mg m − 3 ) Ksat IR MWHC (%) Wetlands Altitude (m) asl cm hr − 1 cm hr − 1 Khorong 2500–2550 1.49 2.81 2.30 47.54 1.30 2.60 2.11 50.04 Tenesolo 2552–2600 1.74 1.38 1.31 41.70 1.52 1.28 1.20 43.90 Khamoqana 2839–2880 1.47 1.56 1.42 43.89 1.28 1.50 1.30 46.20 Khalong-La- Lichelete 2891–2995 1.45 2.84 2.35 44.65 1.26 2.63 2.16 47.00 Lets'eng- La-Likhama 3040–3080 1.45 1.64 1.30 49.23 1.27 1.52 1.19 51.82 Koting-Sa-ha Ramosetsana 3087–3155 1.25 2.92 2.36 54.63 1.09 2.70 2.17 57.51 SE (m)± 0.059 0.120 0.094 1.723 0.016 0.111 0.085 1.814 CD (P < 0.05) 0.019 0.374 0.292 5.368 0.049 0.345 0.266 5.651 CD (P < 0.05) = Critical Difference at less than 5% probability level; SE(m) = Standard Error of the mean; BD= Bulk density, IR= Infiltration rate, Ksat= Saturated hydraulic conductivity, MWHC= Maximum water holding capacity, NS = Non-significant. Table 6 Seasonal variation of physico-chemical properties of the soil as influenced by altitudinal gradients Treatment(s) Spring (Standard wet season) Summer (Peak wet season) CEC (cmol) (P + ) kg − 1 pH EC (dS m − 1 ) SOC (g kg − 1 ) CEC (cmol) (P + ) kg − 1 pH EC (dS m − 1 ) SOC (g kg − 1 ) Wetlands Altitude (m) asl Khorong 2500–2550 28.94 5.97 0.35 83.99 27.09 5.76 0.34 84.67 Tenesolo 2552–2600 25.66 6.16 0.37 68.59 21.50 6.04 0.35 69.14 Khamoqana 2839–2880 28.07 5.93 0.36 72.65 26.45 5.98 0.35 73.24 Khalong-La- Lichelete 2891–2995 27.93 6.22 0.37 79.63 20.86 5.80 0.33 80.27 Lets'eng- La-Likhama 3040–3080 26.10 5.69 0.30 93.59 25.32 5.53 0.29 94.34 Koting-Sa-ha Ramosetsana 3087–3155 31.04 6.19 0.33 95.03 30.11 6.01 0.32 95.80 SE (m)± 1.882 0.074 0.014 0.384 1.939 0.072 0.014 8.28 CD (P < 0.05) NS 0.231 NS 1.197 NS 0.223 NS 18.24 CD (P < 0.05) = Critical Difference at less than 5% probability level; SE(m) = Standard Error of the mean; asl= Above sea level; CEC= Cation exchange capacity; EC= Electrical conductivity; SOC=Soil organic carbon; NS = Non-significant. 4.4. Pearson’s correlation of measured soil attributes during spring (Standard Wet Season) and summer (Peak Wet Season) to evaluate seasonal differences in soil parameter relationships During the standard wet season, sand particles exhibited a significant negative correlation with both silt (− 0.904*) and clay (− 0.884*) (Table 7 a). Clay content showed a significant negative relationship with bulk density (− 0.834*), but was positively correlated with infiltration rate (IR) (0.909*) and soil organic carbon (SOC) (0.812*). Bulk density also displayed a significant negative correlation with IR (− 0.856*). Maximum water holding capacity (MWHC) demonstrated a highly significant positive association with saturated hydraulic conductivity (Ksat) (0.991**). In addition, IR showed a highly significant positive correlation with SOC (0.993**), whereas electrical conductivity (EC) was significantly and negatively correlated with SOC (− 0.845*). During the peak wet season, sand exhibited a significant negative correlation with both silt and clay (Table 7 b). Clay content showed a significant positive correlation with bulk density (BD) (0.855*) and a significant negative correlation with cation exchange capacity (CEC) (− 0.819*). Bulk density was also significantly and negatively correlated with maximum water holding capacity (MWHC) (− 0.885*) and CEC (− 0.934**). MWHC demonstrated a highly significant positive association with soil organic carbon (SOC) (0.932**). Saturated hydraulic conductivity (Ksat) showed a highly significant positive correlation with infiltration rate (IR) (0.993**) and a significant positive correlation with CEC (0.848*). Electrical conductivity (EC) also exhibited a significant negative correlation with SOC (− 0.837*), consistent with the trend observed during the standard wet season. 4.5. Seasonal Index of soil physical, hydraulic and physico-chemical attributes During spring (standard wet season) and summer (peak wet season), several soil properties showed slight deviations from the seasonal average (Figs. 3 , 4 and 5 ). Sand content was 0.2% higher in spring and 0.2% lower in summer relative to the seasonal average (Fig. 3 ). Similarly, silt content was 1% lower in spring but 1% higher in summer. Clay particles were 1.3% above the seasonal average in spring and 1.3% below it in summer. Bulk density was 6.8% higher in spring and correspondingly lower in summer (Fig. 4 ). Saturated hydraulic conductivity showed a 13% increase in spring and a decrease of the same magnitude in summer. Infiltration rate was 5.1% lower in spring and higher in summer. Maximum water holding capacity was 2.6% lower in spring but higher in summer compared with the seasonal average. Table 7 a: Correlation among the soil physical, hydraulic and physico-chemical properties as influenced by altitudinal gradients Sand Sand Silt Clay BD MWHC Ksat IR pH EC CEC SOC 1 Silt -0.904 * 1 Clay -0.884 * 0.600 NS 1 BD 0.627 NS -0.313 NS -0.834 * 1 MWHC 0.073 NS -0.468 NS 0.376 NS -0.642 NS 1 Ksat 0.145 NS -0.526 NS 0.305 NS -0.580 NS 0.991 ** 1 IR -0.736 NS 0.430 NS 0.909 * -0.856 * 0.554 NS 0.470 NS 1 pH 0.251 NS -0.464 NS 0.038 NS 0.011 NS 0.465 NS 0.556 NS -0.084 NS 1 EC 0.719 NS -0.647 NS -0.638 NS 0.491 NS 0.034 NS 0.161 NS -0.712 NS 0.687 NS 1 CEC -0.622 NS 0.639 NS 0.467 NS -0.352 NS -0.028 NS -0.036 NS 0.524 NS -0.125 NS -0.311 NS 1 SOC -0.641 NS 0.355 NS 0.812 * -0.788 NS 0.495 NS 0.383 NS 0.933 ** -0.294 NS -0.845 * 0.266 NS 1 in spring (standard wet season). *Correlation is significant (p < 0.01), **Correlation is highly significant (p < 0.05), BD= Bulk density, IR= Infiltration rate, Ksat= Saturated hydraulic conductivity, EC= Electrical conductivity, CEC= Cation Exchange Capacity and SOC= Soil organic carbon, MWHC= Maximum water holding capacity, NS = Non-significant. Table 7 b: Correlation among the soil physical, hydraulic and physico-chemical properties as influenced by altitudinal gradients in summer (peak wet season). Sand Sand Silt Clay BD MWHC Ksat IR pH EC CEC SOC 1 Silt -0.891 * 1 Clay -0.839 * 0.500 NS 1 BD -0.628 NS 0.285 NS 0.855 * 1 MWHC 0.428 NS -0.099 NS -0.698 NS -0.885 * 1 Ksat 0.400 NS -0.302 NS -0.400 NS -0.668 NS 0.550 NS 1 IR 0.350 NS -0.287 NS -0.323 NS -0.615 NS 0.474 NS 0.993 ** 1 pH 0.206 NS -0.455 NS 0.153 NS 0.000 NS -0.175 NS 0.061 NS 0.138 NS 1 EC -0.146 NS -0.132 NS 0.438 NS 0.612 NS -0.694 NS -0.741 NS -0.678 NS 0.588 NS 1 CEC 0.697 NS -0.426 NS -0.819 * -0.934 ** 0.782 NS 0.848 * 0.800 NS -0.039 NS -0.718 NS 1 SOC 0.274 NS 0.105 NS -0.650 NS -0.777 NS 0.932 ** 0.482 NS 0.392 NS -0.516 NS -0.837 * 0.705 NS 1 Refer to Table 7 a for abbreviations in the footnotes. Soil pH was 1.5% higher in spring and lower in summer, while electrical conductivity was 2.5% higher in spring and lower in summer (Fig. 5 ). Cation exchange capacity was 1.5% lower in spring but higher in summer, and soil organic carbon was 0.4% lower in spring and slightly higher in summer relative to the seasonal average. Discussions 5.1. Seasonal variation of soil particle size distribution, physico-chemical, physical and hydraulic properties across altitudinal gradients The significant variation in soil particle size distribution (PSD) along the altitudinal gradient highlights the important influence of terrain, hydrology, and weathering processes in shaping alpine wetland soils. Soil properties are widely recognized to develop through the interaction of environmental factors such as climate, relief, organisms, parent material, and time, as described in the classical soil-forming factor framework proposed by Hans Jenny. In mountainous environments, elevation modifies climatic variables including temperature and precipitation and also influences vegetation patterns and erosion intensity. These changes affect the mobilization, transport, and deposition of soil particles across landscapes (Liu et al., 2023 ). In the present study, sand content decreased with increasing altitude, while silt and clay fractions increased. This pattern suggests that finer particles tend to accumulate in higher elevation wetlands where runoff energy is relatively low and depositional processes dominate. Conversely, lower elevation sites tend to retain coarser particles because stronger runoff selectively removes finer sediments through erosion. Similar altitudinal patterns have been documented in other mountain ecosystems where enhanced chemical weathering and sediment deposition promote the accumulation of finer soil particles at higher elevations (Nthebere et al., 2025 ). Seasonal variations in PSD further reflect the role of rainfall dynamics in sediment redistribution within wetland environments. The slight increase in sand content and reduction in clay from the standard wet season to the peak wet season may be associated with intensified rainfall and increased runoff during the summer period. Under such conditions, fine particles, particularly clay, are more susceptible to dispersion and transport within saturated soils, whereas sand particles tend to remain relatively stable due to their larger size and mass (Six et al ., 2016; Akor et al., 2025 ). The relatively stable silt fraction indicates that seasonal sediment transport processes mainly affected the finest soil particles. Consequently, the predominance of sandy loam textures at lower elevations and loam textures at higher elevations reflects the long-term interaction between erosion, sediment deposition, and hydrological processes along the altitudinal gradient. The spatial distribution of soil physical and hydraulic properties also corresponded with variations in elevation and soil texture. Bulk density generally decreased with increasing altitude, whereas maximum water holding capacity (MWHC), saturated hydraulic conductivity (Ksat), and infiltration rate (IR) tended to increase at higher elevation wetlands, particularly at Koting-Sa-ha Ramosetsana. Lower bulk density values at higher elevations are commonly associated with greater organic matter accumulation and increased pore space, which reduce soil compaction and improve soil structure (Lal, 2020 ). Wetland soils typically accumulate substantial organic residues because prolonged soil saturation slows microbial decomposition. The accumulation of organic matter promotes aggregate stability and the formation of interconnected pore networks, thereby enhancing water retention and hydraulic conductivity (Reddy and DeLaune, 2017). Consequently, the higher MWHC, Ksat, and IR recorded at upper elevation wetlands likely reflect improved soil structure and increased porosity associated with organic-rich soils. Seasonal differences in these physical and hydraulic parameters further illustrate the influence of soil moisture dynamics. Higher bulk density, infiltration rate, and hydraulic conductivity during the standard wet season may indicate relatively stable soil structure before prolonged saturation occurs. As rainfall intensifies during the peak wet season, extended waterlogging may lead to swelling of fine particles and partial blockage of soil pores, thereby reducing infiltration and hydraulic conductivity (Lal, 2020 ). In contrast, maximum water holding capacity increased during the peak wet season, reflecting the enhanced moisture retention capacity of soils under saturated conditions. These responses highlight the close interaction between hydrological processes and soil physical functioning in wetland ecosystems. The chemical characteristics of the soils also exhibited patterns associated with elevation and seasonal dynamics. Soil pH values remained within the slightly acidic range across the wetlands and varied significantly along the altitudinal gradient. Lower pH values at higher elevations are commonly associated with increased rainfall and enhanced leaching of base cations such as calcium, magnesium, potassium, and sodium (Lal, 2020 ). Furthermore, the decomposition of organic residues in wetland soils produces organic acids that can contribute to soil acidification. Electrical conductivity (EC) and cation exchange capacity (CEC), however, did not show strong altitudinal responses, suggesting that these properties may be more strongly influenced by soil mineral composition and organic matter content than by elevation alone. The relatively low EC values observed across the wetlands indicate minimal salt accumulation, which is typical of environments characterized by high precipitation and continuous water movement (Reddy and DeLaune, 2017). Soil organic carbon (SOC) exhibited a clear increase with altitude, with the highest concentrations observed at Koting-Sa-ha Ramosetsana and the lowest at Tenesolo. This trend is consistent with ecological processes commonly observed in mountainous environments where cooler temperatures and higher moisture levels reduce microbial activity and slow organic matter decomposition (Blagodatskaya et al., 2016 ; Lal, 2020 ). Wetland conditions further intensify this effect because saturated soils create anaerobic environments that limit microbial mineralization of organic matter (Reddy and DeLaune, 2017). As a result, organic carbon accumulates more readily over time. Elevated SOC levels at higher elevation wetlands therefore reflect the combined influence of cooler climatic conditions, persistent soil moisture, and slower decomposition rates. In addition to reflecting biogeochemical processes, SOC is widely recognized as an important indicator of wetland ecological health. High SOC concentrations often indicate strong carbon sequestration capacity, sustained vegetation productivity, and stable hydrological conditions (Mitsch et al., 2015 ). The relatively high SOC values observed at the upper elevation wetlands therefore suggest that these sites may possess comparatively better ecological condition and greater carbon storage potential. Seasonal variations in soil chemical properties were less pronounced than the altitudinal differences. Slight decreases in EC, CEC, and pH during the peak wet season may be attributed to increased leaching under conditions of intense rainfall and prolonged soil saturation. Conversely, SOC showed a modest increase during the peak wet season, likely reflecting enhanced plant growth and greater organic residue inputs during the summer growing period combined with slower decomposition under waterlogged conditions (Lal, 2020 ). Although the results revealed clear altitudinal and seasonal patterns, several limitations should be considered. Soil sampling was conducted only during the standard wet season and the peak wet season, and therefore does not capture soil dynamics during the dry season when moisture depletion, oxidation of organic matter, and soil compaction may significantly alter soil characteristics. In addition, variations in water table depth and short-term rainfall intensity, important drivers of wetland soil processes, were not directly measured. Future studies incorporating year-round monitoring and detailed hydrological measurements would provide a more comprehensive understanding of how climatic variability and elevation interact to influence soil processes in alpine wetlands. Overall, the findings demonstrate that both elevation and seasonal hydrological conditions play critical roles in shaping the physical, hydraulic, and chemical properties of alpine wetland soils. Variations in soil texture, bulk density, water retention capacity, and organic carbon storage reflect the underlying soil-forming processes operating within mountain wetland landscapes and provide valuable insight into the functioning and ecological condition of these sensitive ecosystems. 5.2. Seasonal Index of soil physical, hydraulic and physico-chemical attributes and Pearson’s correlation of measured soil attributes during late spring and summer The correlation patterns observed between soil properties during the standard wet season (spring) and the peak wet season (summer) reflect the dynamic hydrological and structural processes that regulate soil functioning in alpine wetlands. The strong negative correlation of sand with both silt and clay in both seasons indicates the inherent textural trade-off within soil particle size distribution, where an increase in coarse particles occurs at the expense of finer fractions. Such relationships are typical in wetland soils and influence water retention, infiltration, and nutrient dynamics along altitudinal gradients (Zhao et al., 2023 ; Li et al ., 2020). During the standard wet season, clay showed a significant negative relationship with bulk density but positive correlations with infiltration rate and soil organic carbon. This pattern suggests that finer particles combined with organic matter promote the formation of stable soil aggregates and pore networks, which improve infiltration and reduce soil compaction. In alpine wetlands, higher SOC associated with clay fractions enhances soil structure and water transmission, thereby supporting wetland hydrological regulation and ecosystem functioning (Minasny and McBratney, 2018). The negative correlation between bulk density and infiltration rate further confirms that compacted soils restrict water movement, while less dense soils allow rapid infiltration and groundwater recharge. The highly significant association between maximum water holding capacity and saturated hydraulic conductivity suggests that soils with greater pore space can simultaneously retain and transmit water efficiently. This is particularly important in alpine wetlands, where seasonal water storage regulates downstream hydrology and maintains wetland resilience during fluctuating moisture conditions (Hribljan et al ., 2016). The strong positive correlation between infiltration rate and SOC also highlights the ecological role of organic matter in improving soil porosity and permeability. Conversely, the negative correlation between electrical conductivity and SOC suggests that higher organic matter may dilute soluble salts or promote leaching, reducing salinity levels within wetland soils (Yang and Guo, 2019 ). During the peak wet season, several correlations shifted slightly, indicating seasonal hydrological influence. The positive relationship between clay and bulk density suggests temporary compaction of fine particles under prolonged saturation. At the same time, the negative correlation between clay and cation exchange capacity may reflect dilution or redistribution of exchangeable ions under high moisture conditions. The strong negative relationships of bulk density with both maximum water holding capacity and CEC emphasize that compact soils limit water retention and nutrient exchange processes, which are critical for wetland productivity. Hydraulic parameters also showed strong seasonal coupling. The highly significant positive relationship between saturated hydraulic conductivity and infiltration rate indicates synchronized water movement through soil pores during peak rainfall periods. Additionally, the positive association of saturated hydraulic conductivity with CEC suggests that soils capable of transmitting water efficiently may also facilitate nutrient mobility and exchange. The slight seasonal deviations in particle size distribution and soil physical properties indicate that alpine wetlands respond sensitively to seasonal hydrological cycles. Lower bulk density and higher infiltration and water holding capacity in summer reflect improved soil structure under sustained moisture conditions. Collectively, these correlations demonstrate that interactions among soil texture, organic carbon, and hydraulic properties regulate water storage, nutrient retention, and ecological stability in alpine wetland systems. Implications and policy recommendation The results highlight several implications for the effective conservation and management of alpine wetlands in Lesotho. Variations in soil texture, hydraulic behaviour, and soil organic carbon (SOC) along the altitudinal gradient suggest that wetlands located at higher elevations play a crucial ecological role as natural reservoirs for water storage and as significant carbon sinks. These functions contribute to regulating hydrological processes and maintaining ecological balance in mountainous landscapes. Nevertheless, the presence of fine soil particles that can easily be displaced during intense rainfall or surface disturbance indicates that these wetlands remain highly sensitive to external pressures such as excessive grazing, livestock trampling, and poorly managed land-use activities. Consequently, environmental policies should place greater emphasis on safeguarding high-altitude wetlands where elevated SOC levels and improved soil structure reflect relatively healthier ecosystem conditions and stronger carbon storage capacity. Practical management measures may include regulating grazing intensity, rehabilitating degraded wetland areas, and creating protective buffer zones around vulnerable wetland systems to minimize soil compaction and sediment displacement. Furthermore, incorporating systematic wetland monitoring into national environmental management programmes would enable continuous evaluation of soil quality and hydrological performance. Considering the strong influence of seasonal rainfall variability on soil processes, adaptive land management approaches that respond to climatic fluctuations are equally important. Encouraging community participation in wetland stewardship and promoting sustainable rangeland practices can strengthen the long-term resilience of alpine wetlands while sustaining essential ecosystem services such as water regulation, biodiversity conservation, and carbon retention within Lesotho’s highland regions. Conclusion In conclusion, the results indicated that both elevation and seasonal hydrological conditions play a major role in shaping soil texture, hydraulic characteristics, and chemical properties in alpine wetlands. The higher altitude wetland, Koting-Sa-ha Ramosetsana, contained finer soil fractions, lower bulk density, greater water retention capacity, and higher soil organic carbon compared with Tenesolo, suggesting better soil structure, stronger carbon storage potential, and improved ecological status. Seasonal rainfall patterns also influenced sediment movement and soil water dynamics. These outcomes highlight the hydrological and ecological significance of these alpine wetlands. Future studies should incorporate continuous annual monitoring, vegetation assessments, and water-table measurements to improve understanding of long-term soil and wetland processes. Declarations CRediT authorship contribution statement Knight Nthebere and Mosiuoa Mochala: Writing – review and editing, Writing – original draft. Knight Nthebere: Conceptualization. Knight Nthebere and Mosiuoa Mochala: Formal analysis. Knight Nthebere and Mosiuoa Mochala: Methodology and Data curation. Knight Nthebere: Investigation. Funding The research project was funded by the Deutsche Gesellschaft für Internationale Zusammenarbeit (GIZ) and the German Government. The funds were managed by the WaterNet Trust. Data Availability Statement Data are provided within the manuscript or Supplementary Information files. Acknowledgments The authors are extremely thankful to the Government of Lesotho, in general, and the ReNOKA movement in particular, which initiated and coordinated the implementation of the Integrated Catchment Management Programme in Lesotho, the EU, Deutsche Gesellschaft für Internationale Zusammenarbeit (GIZ) and the German Government for providing funds for the implementation of this study, as well as WaterNet for managing the research fund. Conflicts of Interest The authors declare that they have no competing interests. References Akor S, Flores AN, Rudisill W, Bergstrom A, McNamara JP (2025) Impact of cloud microphysics schemes and boundary conditions on modeled snowpack in mountain watersheds. 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Water 15(6):1140 Supplementary Materials Supplementary Fig 1: Overview of Tenesolo (a), Khorong (b), Khamoqana (c), Khalong-La- Lichete (d), Lets’eng-La-Likhama (e) and Koting-Sa- ha Ramosetsana (f). Lon= Longitude, Lat= Latitude Supplementary Table S. Ecological class utilized for percentage score (PES) evaluations of inland aquatic ecosystems in South Africa, along with the applicable range of PESs for each class (Kleynhans, Macfarlane et al (1996) 2009) [color coding is per the River Eco-Status Monitoring Programme (REMP) of DWS] Additional Declarations The authors declare no competing interests. Supplementary Files SupplementaryMaterials.docx Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9212062","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":611397798,"identity":"05ae9c98-cbf1-431f-89f1-8d230cbfd228","order_by":0,"name":"Knight Nthebere","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2ElEQVRIiWNgGAWjYBACCQYGNgbGBiCLHUQYWBCjhRmqhecASIsEKVokEqC2EgKSs/uPPfi4wyafX/L51Q0/CiQY+Nu7E/BqkZY5zG4480ya5czZOWU3e4AOkzhzdgNeLXISyWzSvG2HDQxu56Td4AFqMZDIJULL37b/BvY3z6Td/EOMFmmQFsa2AwYGEuzHbhNli+SMZDPJ3jPJBhJncthuyxhI8BD0i8SNxGcSP3fYGfC3H392880fGzn+9l78WpAAjwGYJFY5CLA/IEX1KBgFo2AUjCAAABbyQ38EgeDtAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0002-9933-1365","institution":"National University of Lesotho","correspondingAuthor":true,"prefix":"","firstName":"Knight","middleName":"","lastName":"Nthebere","suffix":""},{"id":611397799,"identity":"573be5ac-8dd8-4ec7-b054-f4f40466b121","order_by":1,"name":"Mosiuoa Mochala","email":"","orcid":"","institution":"National University of Lesotho","correspondingAuthor":false,"prefix":"","firstName":"Mosiuoa","middleName":"","lastName":"Mochala","suffix":""}],"badges":[],"createdAt":"2026-03-24 12:44:03","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":true,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":true},"doi":"10.21203/rs.3.rs-9212062/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9212062/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":105368197,"identity":"64b1f73f-d7e2-4def-9511-eaad1f4ccf26","added_by":"auto","created_at":"2026-03-25 08:57:56","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":594462,"visible":true,"origin":"","legend":"\u003cp\u003eMap illustrating the location of Lesotho within Africa, showing all ten administrative districts and the selected alpine wetland study areas. The highlighted districts; Mokhotlong and Thaba-Tseka, represent the regions where the research was conducted. Black dots mark the positions of the identified alpine wetlands\u003c/p\u003e","description":"","filename":"image1.png","url":"https://assets-eu.researchsquare.com/files/rs-9212062/v1/9387fe03a7169077db80e48e.png"},{"id":105368195,"identity":"194105f1-94ee-4778-8231-d812b94f8ffb","added_by":"auto","created_at":"2026-03-25 08:57:56","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":455083,"visible":true,"origin":"","legend":"\u003cp\u003eA GPS-based map illustrating the Upper Senqu main catchment, encompassing the three sub-catchments; Khubelu, Senqunyane, and Sani, together with the associated wetland locations. The spatial representation of the map was produced using ArcGIS software (Version 3.5.x).\u003c/p\u003e","description":"","filename":"image2.png","url":"https://assets-eu.researchsquare.com/files/rs-9212062/v1/955670781eb169abd4e38443.png"},{"id":105368183,"identity":"5bd2603e-4287-4df7-808d-ce36b00a3a77","added_by":"auto","created_at":"2026-03-25 08:57:52","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":28709,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSeasonal index of soil particle size distribution as influenced by altitudinal gradients\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-9212062/v1/91d64f527906ee17511fa440.png"},{"id":105368189,"identity":"402192e6-2194-45bd-9c00-25d747ee86e4","added_by":"auto","created_at":"2026-03-25 08:57:53","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":34429,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSeasonal index of soil physical and hydraulic properties as influenced by altitudinal gradients. BD= Bulk density, IR= Infiltration rate, Ksat= Saturated hydraulic conductivity and MWHC= Maximum water holding capacity.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-9212062/v1/00e519c20af67edd5925d6d4.png"},{"id":105368214,"identity":"6ff02e50-d7e6-4678-a1c4-7b21bfe13ff3","added_by":"auto","created_at":"2026-03-25 08:57:58","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":31943,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSeasonal index of soil physico-chemical properties as influenced by altitudinal gradients. EC= Electrical conductivity, CEC= Cation exchange capacity and SOC= Soil organic carbon.\u003c/strong\u003e\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-9212062/v1/37be4f114aba26f8580f84dd.png"},{"id":105368240,"identity":"fa99cd99-c9e1-4924-a4a6-d7779d8aa767","added_by":"auto","created_at":"2026-03-25 08:58:10","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3196233,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9212062/v1/13b8d517-3361-4b12-b9bf-721e12b9c21b.pdf"},{"id":105368185,"identity":"098b0bad-e85e-4edb-8e1d-5602d4923e78","added_by":"auto","created_at":"2026-03-25 08:57:53","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":1220477,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryMaterials.docx","url":"https://assets-eu.researchsquare.com/files/rs-9212062/v1/b9391f148f3669454ebb4b95.docx"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003e\u003cstrong\u003eSeasonal Dynamics of Physical, Hydraulic, and Physico-Chemical Attributes of the Soil across Altitudinal Gradients in the Alpine Wetlands of Lesotho\u003c/strong\u003e\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eAlpine wetlands are increasingly acknowledged as ecologically fragile yet functionally indispensable components of mountain environments, where they modulate hydrological fluxes, conserve biodiversity, and attenuate climate variability (IPCC, 2021; Mitsch et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Within the high-elevation landscapes of Lesotho, these wetlands constitute critical headwater systems that regulate stream discharge, facilitate groundwater recharge, and safeguard downstream water supplies, including key transboundary water resources. Emerging evidence indicates that, despite occupying relatively small spatial extents, mountain wetlands exert disproportionately large influences on watershed hydrology and carbon sequestration (Lin et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Nthebere et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The ecological performance of alpine wetlands is fundamentally regulated by soil physical, hydraulic, and physico-chemical characteristics, which collectively influence water retention, infiltration dynamics, aeration, nutrient turnover, and carbon stabilization. Consequently, examining the temporal variability of these soil attributes, particularly across seasons, is essential for advancing sustainable wetland management and enhancing climate resilience in highland systems (Nthebere et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSoil physical properties, such as bulk density, total porosity, texture, and aggregate stability, determine structural coherence, resistance to erosion, and root penetration. Hydraulic attributes, including infiltration rate, saturated hydraulic conductivity, and maximum water holding capacity, regulate soil-water interactions, influencing runoff generation and aquifer recharge. Key physico-chemical indicators, notably soil pH, electrical conductivity and soil organic carbon (SOC), govern microbial activity and biogeochemical cycling that sustain ecosystem productivity (Lal, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Lehmann et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). These interrelated properties are dynamic, varying with seasons, vegetation across altitude (Mirza and Patil, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Elevation-driven changes in plant communities significantly influence SOC stabilization and soil hydraulic behavior (Zhang \u003cem\u003eet al\u003c/em\u003e., 2022). Soil characteristics are further shaped by interactions among abiotic factors; temperature, precipitation, topography and biotic components, including plants and microbes, which fluctuate spatially and seasonally (Das et al, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Liu et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeasonal transitions, particularly from the standard wet season to the peak wet season, modify soil moisture, redox status, hydraulic conductivity, and nutrient availability. Progressive wetting may enhance aggregate stability and microbial mineralization, whereas sustained saturation can reduce aeration, destabilize soil structure, limit infiltration, and promote denitrification and methane production (Lin et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Altitudinal gradients mediate these effects by influencing thermal regimes, rainfall intensity, vegetation composition, and decomposition dynamics, yet these interactions remain poorly documented in southern African alpine wetlands. Despite their ecological and hydrological importance, Lesotho\u0026rsquo;s alpine wetlands are rarely studied with integrated assessments that combine seasonal variability, altitudinal differentiation, and soil physical, hydraulic, and physico-chemical indicators. Most studies focus on single properties or seasons, overlooking interactive effects critical under increasing climate variability and anthropogenic pressures such as overgrazing and land-use intensification (IPCC, 2021). This study addresses the central question: How do seasonal transitions from the standard to peak wet season, across altitudinal gradients, influence the integrated soil functionality of Lesotho\u0026rsquo;s alpine wetlands? It is hypothesized that seasonal progression significantly alters soil physical, hydraulic, and physico-chemical properties, with systematic variation along elevation gradients.\u003c/p\u003e \u003cp\u003eThe study aimed to quantify seasonal changes in soil attributes across elevation zones during both wet-season phases and to evaluate a composite seasonal index of selected soil properties reflecting functional shifts along altitudinal gradients. Assessing soil variability via an integrated index is vital because alpine wetlands regulate watershed hydrology, influence SOC stabilization and greenhouse gas fluxes, and provide a quantitative framework to detect changes that individual indicators may miss. Integrating seasonal and altitudinal controls enhances predictive capacity under climate change and supports sustainable wetland management in Lesotho\u0026rsquo;s mountainous landscapes. By combining seasonal dynamics with elevation-driven controls through a holistic soil quality framework, this study offers a comprehensive approach to understanding soil functionality and ecological resilience in alpine wetlands, informing climate adaptation and watershed management strategies.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Description of the Study Area\u003c/h2\u003e \u003cp\u003eThe investigation was conducted in the high-altitude regions of Lesotho, spanning elevations between 2500 and 3089 m above sea level within the mountain agro-ecological zones (AEZs) of Mokhotlong and Ha-Mohale. The study focused on three sub-catchments; Khubelu, Senqunyane, and Sani, situated within the Upper Senqu main catchment. Two alpine wetlands were identified and sampled in each sub-catchment, giving a total of six study wetlands. In the Khubelu sub-catchment, the Lets\u0026rsquo;eng-la-Likhama and Koting-sa-ha Ramosetsana wetlands were included. The Senqunyane sub-catchment comprised the Khorong and Tenesolo wetlands, while the Sani sub-catchment encompassed the Khamoqana and Khalong-la-Lichelete wetlands. Figure\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e presents a map of Lesotho, indicating its ten administrative districts within the African continent and highlighting the selected alpine wetland sites. Figure\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e illustrates the GPS-based layout of the Upper Senqu main catchment, showing the location of the three sub-catchments; Khubelu, Senqunyane, and Sani, and the distribution of the sampled wetlands within them.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 The Alpine Wetland Features\u003c/h2\u003e \u003cp\u003eAcross the alpine wetlands, human settlement is sparse, with activity largely confined to seasonal cattle posts. Within Lesotho\u0026rsquo;s highland agro-ecological zones, vegetation is predominantly composed of shrubs and grasses adapted to high-altitude conditions. The underlying geology of these wetlands is mainly associated with formations described as Lesotho Genesis (Schmitz and Rooyani, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1987\u003c/span\u003e). At Tenesolo, the wetland showed pronounced degradation, largely attributable to repeated vegetation burning, past overgrazing, and extensive burrowing by ice rats (Supplementary Figure S1a). In contrast, land use around Khorong was primarily characterized by cultivation (Supplementary Figure S1b). In Khamoqana, burrowing by ice rats resulted in numerous surface holes, accelerated soil loss through runoff, and the dominance of sparse vegetation with shallow root systems (Supplementary Figure S1c). Khalong-La-Lichelete was distinguished by the occurrence of deep-rooted shrub species (Supplementary Figure S1d). The Lets\u0026rsquo;eng-La-Likhama wetland was mainly affected by limited grass cover, erosion linked to inadequate vegetative protection, and heavy grazing pressure from sheep (Supplementary Figure S1e). At Koting-Saha Ramosetsana, shrubs predominated in the surrounding landscape, while grasses were more common within the wetland itself (Supplementary Figure S1f).\u003c/p\u003e \u003cp\u003eSoil degradation across all sites was assessed using WET-Health version 2.0, an advanced assessment framework developed to evaluate the current ecological status of wetland systems, following the approaches outlined by Kleynhans (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1996\u003c/span\u003e) and Macfarlane et al. (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). The assessment outcomes are presented in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, with additional detail provided in Supplementary Table S1.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eAlpine wetland characteristics.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eAlpine Wetlands\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eLatitude S\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eLongitude E\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eSoil Degradation Level Assessed with WET Health According to Kleynhans (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1996\u003c/span\u003e) and Macfarlane et al. (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2009\u003c/span\u003e).\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePES Score (%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eDescription\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhorong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u0026minus;29.457168\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28.268082\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLargely natural with few modifications. A slight change in ecosystem processes is discernible, and a small loss of natural habitats and biota may have taken place.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTenesolo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u0026minus;29.449256\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28.149214\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eExtensively altered and alterations in ecological functions accompanied by the disappearance of natural habitats and native species.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhamoqana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u0026minus;29.457178\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28.268094\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSeriously modified, the change in ecosystem processes, great loss of natural habitat and biota but some remaining natural habitat features are still being recognized.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhalong-La-Lichelete\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u0026minus;29.563552\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e29.247207\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUnmodified natural wetland\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLets\u0026rsquo;eng-La-Likhama\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u0026minus;29.076355\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28.836095\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLargely modified with a large change in ecosystem processes and loss of natural habitat, and biota has occurred.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKoting-Sa-ha Ramosetsana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e\u0026minus;29.022686\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28.871324\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLargely natural with few modifications and a slight change in ecosystem processes being discernible and a small loss of natural habitats and biota may have taken place.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eS\u0026thinsp;=\u0026thinsp;Southern Hemisphere; E\u0026thinsp;=\u0026thinsp;Eastern Hemisphere; PES\u0026thinsp;=\u0026thinsp;Percentage.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3. Climate\u003c/h2\u003e \u003cp\u003eLesotho\u0026rsquo;s climatic conditions are largely influenced by its position on the central southern African Plateau. The country experiences a sub-humid, cool temperate climate, with distinct seasonal contrasts marked by hot, wet summers and cold, dry winters (Wu et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). June is generally the coldest month, when mean minimum temperatures are close to 0\u0026deg;C. In the lowland zones, winter temperatures commonly range from about \u0026minus;\u0026thinsp;1\u0026deg;C to \u0026minus;\u0026thinsp;3\u0026deg;C, while the high-altitude regions may experience more severe conditions, with temperatures declining to between roughly \u0026minus;\u0026thinsp;6\u0026deg;C and \u0026minus;\u0026thinsp;8.5\u0026deg;C (Malebajoa, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Average annual temperatures are approximately 15.2\u0026deg;C in the lowlands and about 7\u0026deg;C in the highlands.\u003c/p\u003e \u003cp\u003eJanuary typically registers the highest average maximum temperatures, reaching nearly 32\u0026deg;C in the lowlands and around 20\u0026deg;C in the highlands. Rainfall distribution varies geographically, generally ranging from about 500 mm to 1200 mm per year, with the northern and eastern parts of the country receiving comparatively higher amounts. The majority of precipitation, around 85%, occurs during the summer months from October to April. Frost and occasional snowfall are common features of winter in the mountainous areas. Mean annual rainfall in the Tenesolo and Khorong wetlands is close to 1000 mm, while slightly higher averages of approximately 1044 mm are reported for the Khamoqana, Khalong-La-Lichelete, Lets\u0026rsquo;eng-La-Likhama, and Koting-Sa-ha Ramosetsana wetlands (Malebajoa, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.4. Research Design\u003c/h2\u003e \u003cp\u003eA block experimental layout was adopted, whereby differences in altitude among alpine wetlands were organized into blocks defined by catchments sharing similar attributes, particularly wetland condition (degraded versus intact) and elevation span, in order to reduce extraneous variation. Controlling for these site characteristics allowed clearer evaluation of the treatment effect, represented by altitudinal differences among wetlands. Altitude was therefore considered the main experimental factor and was represented by six alpine wetlands distributed across three sub-catchments; Khubelu, Senqunyane, and Sani, with two wetlands selected from each catchment. Specific treatment descriptions, corresponding to the altitudinal classes of the selected wetlands, are presented in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e2.5. Wetland Selection Criteria\u003c/h2\u003e \u003cp\u003eWetland sites were selected based on specific elevation thresholds within each catchment: \u0026ge;3000 m in Khubelu, \u0026ge;\u0026thinsp;2500 m in Senqunyane, and \u0026ge;\u0026thinsp;2800 m in Sani. For every catchment, two alpine wetlands were purposively selected to reflect contrasting ecological states, one exhibiting signs of degradation and the other remaining in a comparatively sound condition. The health status of the wetlands was evaluated using the WET-Health assessment tool (version 2.0) (Kleynhans, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Macfarlane et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2009\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e2.6. Selection of Sampling Points\u003c/h2\u003e \u003cp\u003eSampling locations within each wetland were determined using a stratified random approach to ensure adequate representation of site variability. Within each stratum, a grid-based layout was applied to systematically position sampling points in alignment with the soil sampling objectives (Paulsen et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). To allow replication, every wetland was divided into four equal grid sections, yielding four replicate samples per site.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eTreatment details.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"2\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eTreatment(s)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAlpine Wetlands\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAltitude (m) asl\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhorong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2500\u0026ndash;2550\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTenesolo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2552\u0026ndash;2600\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhamoqana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2839\u0026ndash;2880\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhalong-La-Lichelete\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2891\u0026ndash;2995\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLets\u0026rsquo;eng-La-Likhama\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3040\u0026ndash;3080\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKoting-Sa-ha Ramosetsana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3087\u0026ndash;3155\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e2.7. Soil Sampling and Standard Analytical Methods\u003c/h2\u003e \u003cp\u003eSoil samples were collected at the end of September 2024 (late spring), representing the standard wet season, and again in February 2025 (summer), corresponding to the peak wet season. These two sampling periods were deliberately selected to capture intra-seasonal variability and to facilitate the development of a seasonal index reflecting shifts in soil physical, hydraulic, and physico-chemical properties under varying moisture regimes in alpine wetlands. Sampling during both phases of the wet season enabled comparison between early-season conditions, when soils begin to re-wet, and peak-season conditions, when moisture availability and hydrological activity are typically at their maximum.\u003c/p\u003e \u003cp\u003eAt each wetland, soil was collected from the 0\u0026ndash;15 cm depth, representing the active surface layer most responsive to seasonal changes and biogeochemical processes. Within each replication, ten subsamples were taken from multiple points and thoroughly mixed to form a single composite sample. Four replications were obtained per site. The composite samples were air-dried under shade, lightly crushed, and passed through 2.0 mm and 0.5 mm sieves to obtain the required fractions for analysis. After labeling, the processed samples were stored in polyethylene bags pending laboratory evaluation. Physical, hydraulic, and physico-chemical analyses were subsequently conducted using established standard procedures (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eMethodology and terms of the references adopted for the analysis of physical, hydraulic and physico-chemical properties of the soil at the 0\u0026ndash;15 cm depth.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eS.No\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSoil Property\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMethod\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eReference\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eMechanical separates\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003eHydrometer method\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" morerows=\"3\" nameend=\"c5\" namest=\"c4\" rowspan=\"4\"\u003e \u003cp\u003eGee and Or (2002)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSand (%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSilt (%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eClay (%)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSoil reaction (pH)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSoil: water suspension (1:2.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" morerows=\"1\" nameend=\"c5\" namest=\"c4\" rowspan=\"2\"\u003e \u003cp\u003eJackson (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1973\u003c/span\u003e)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eElectrical conductivity (dS m\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBulk density (Mg m\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003eCore sampler\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBlake and Hartge (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1986\u003c/span\u003e)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSoil organic carbon (g kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003eWet oxidation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eWalkley and Black (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1934\u003c/span\u003e)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCation exchange capacity (cmol) (P\u003csup\u003e+\u003c/sup\u003e) kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003eSodium acetate method\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBower et al. (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1952\u003c/span\u003e)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSaturated hydraulic conductivity (cm hr\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003eConstant head method\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eKlute and Driksen (1986)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eInfiltration rate (cm hr\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c4\" namest=\"c3\"\u003e \u003cp\u003eDouble-ring infiltrometer\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBouwer (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1986\u003c/span\u003e)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMaximum water holding capacity (%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eKeen\u0026rsquo;s cup method\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eKeen-Raczkowski (1921)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Statistical Analysis","content":"\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe collected data were subjected to statistical evaluation using analysis of variance (ANOVA) based on a one\u0026ndash;factor randomized block design, following the procedure outlined by Panse and Sukhatme (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1978\u003c/span\u003e). Differences among treatment means were tested for statistical significance at the 5% probability level. Relationships among the measured soil attributes were further examined using Pearson\u0026rsquo;s correlation analysis and principal component analysis (PCA). These analyses were conducted using the SQI CAL tool (Mohanty, 2020), which is specifically developed for soil quality assessment. The analyses were performed independently for each season in order to evaluate seasonal differences in the relationships among soil parameters. Several techniques are available for determining seasonal indices; however, the simple average method was adopted in this study as described by Mirza and Patil (\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Seasonal indices were computed to quantify temporal variability in the measured soil properties. In this context, the seasonal index (SI) for spring (representing the standard wet season) and summer (representing the peak wet season) was calculated to express the average percentage deviation of soil parameters during each season relative to the overall mean.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e3.1. Method of simple averages\u003c/h2\u003e \u003cp\u003eA simple averaging approach was applied to assess seasonal variations over the time series. The seasonal index (S.I) was computed by expressing the mean value for each period as a percentage of the overall mean x \u003cem\u003ei.e\u003c/em\u003e., seasonal index for different periods (Mirza and Patil, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e);\u003c/p\u003e \u003cp\u003eS.I\u0026thinsp;=\u0026thinsp;Average of a season / Grand Average of the all seasonal averages \u0026times;100\u003c/p\u003e \u003cp\u003eSeasonal index for i\u003csup\u003eth\u003c/sup\u003e season,\u003c/p\u003e \u003cp\u003eSi\u0026thinsp;=\u0026thinsp;Average of i\u003csup\u003eth\u003c/sup\u003e season / Grand average of k seasons \u0026times; 100\u003c/p\u003e \u003cp\u003eTherefore, Si\u0026thinsp;=\u0026thinsp;x\u003csub\u003e1\u003c/sub\u003e / x \u0026times; 100; i\u0026thinsp;=\u0026thinsp;1, 2, 3\u0026hellip; k\u003c/p\u003e \u003cp\u003eThus seasonal indices are,\u003c/p\u003e \u003cp\u003ex\u003csub\u003e1\u003c/sub\u003e /x \u0026times; 100, x\u003csub\u003e2\u003c/sub\u003e / x \u0026times; 100,\u0026hellip;.., x k/ x \u0026times; 100.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e4.1. Seasonal variation of soil particle size distribution and texture across altitudinal gradients\u003c/h2\u003e \u003cp\u003eSoil particle size distribution (PSD) differed significantly between the standard wet and peak wet seasons along the altitudinal gradients of the selected alpine wetlands (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Across both seasons, sand, silt and clay contents ranged from 39.79\u0026ndash;64.97%, 21.92\u0026ndash;35.72% and 10.45\u0026ndash;25.29%, respectively. Sand content showed a clear decline with increasing altitude, with significantly higher proportions recorded at Khorong, Tenesolo and Khalong-La-Lichelete, while lower values occurred at Lets'eng-La-Likhama and Koting-sa-ha Ramosetsana. In contrast, silt and clay fractions generally increased from Tenesolo to Koting-sa-ha Ramosetsana along the elevation gradient. Seasonal comparison indicated a slight increase in sand content from the standard wet season (late spring) to the peak wet season (summer), whereas clay content declined during the same period, while silt remained relatively stable. Consequently, soils at Khorong, Tenesolo and Khalong-La-Lichelete were classified as sandy loam, whereas those at Khamoqana, Lets'eng-La-Likhama and Koting-sa-ha Ramosetsana were predominantly loam in both the seasons (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSeasonal variation of soil particle size distribution and texture as influenced by altitudinal gradients\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" morerows=\"1\" nameend=\"c2\" namest=\"c1\" rowspan=\"2\"\u003e \u003cp\u003eTreatment(s)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c5\" namest=\"c3\"\u003e \u003cp\u003eSpring (Standard wet season)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e \u003cp\u003eSummer (Peak wet season)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eTextural Class\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSand\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSilt\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eClay\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSand\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSilt\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eClay\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWetlands\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAltitude (m) asl\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"6\" nameend=\"c8\" namest=\"c3\"\u003e \u003cp\u003e(%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhorong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2500\u0026ndash;2550\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e64.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e23.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e11.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e64.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e23.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e11.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSandy loam\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTenesolo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2552\u0026ndash;2600\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e60.48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e28.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e10.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e60.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e28.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e10.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSandy loam\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhamoqana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2839\u0026ndash;2880\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e52.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e34.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e52.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e34.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e13.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eLoam\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhalong-La- Lichelete\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2891\u0026ndash;2995\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e64.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e21.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e64.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e21.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e13.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSandy loam\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLets'eng- La-Likhama\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3040\u0026ndash;3080\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e46.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e35.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e18.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e46.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e35.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e17.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eLoam\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKoting-Sa-ha Ramosetsana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3087\u0026ndash;3155\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e39.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e35.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e25.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e39.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e35.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e25.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eLoam\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSE (m)\u0026plusmn;\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.079\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.004\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.082\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.279\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.140\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.245\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eCD (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.246\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.013\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.257\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.868\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.307\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.764\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"9\"\u003eCD (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) = Critical Difference at less than 5% probability level; SE(m) = Standard Error of the mean\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003e4.2. Seasonal variation of soil physical and hydraulic properties across altitudinal gradients\u003c/h2\u003e \u003cp\u003eThe soil physical property (bulk density) and hydraulic attributes; maximum water holding capacity, saturated hydraulic conductivity, and infiltration rate, showed significant seasonal variation along the altitudinal gradients (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Across the wetlands, bulk density (BD), maximum water holding capacity (MWHC), saturated hydraulic conductivity (Ksat), and infiltration rate (IR) ranged from 1.09\u0026ndash;1.74 Mg m⁻\u0026sup3;, 41.70\u0026ndash;57.51%, 1.28\u0026ndash;2.92 cm hr⁻\u0026sup1;, and 1.20\u0026ndash;2.36 cm hr⁻\u0026sup1;, respectively. Bulk density declined with increasing altitude and was significantly lower at Koting-Sa-ha Ramosetsana (KSHM), recording 1.25 Mg m⁻\u0026sup3; during the standard wet season and 1.09 Mg m⁻\u0026sup3; during the peak wet season. In both seasons, MWHC, Ksat, and IR were significantly higher at KSHM, whereas the lowest values (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) were observed at Tenesolo compared with the other wetlands along the altitudinal gradient (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Seasonal trends indicated that BD, Ksat, and IR were higher during the standard wet season but declined during the peak wet season, while MWHC increased from the standard wet season to the peak wet season across all wetlands.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e4.3. Seasonal variation of soil physico-chemical properties across altitudinal gradients\u003c/h2\u003e \u003cp\u003eSoil pH was significantly influenced by altitudinal gradients across the alpine wetlands, ranging from 5.53\u0026ndash; 6.22 in both seasons (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). The lowest and highest pH values were recorded at Lets\u0026rsquo;eng-la-Likhama (LLL) and Khorong, respectively, indicating slightly acidic soil conditions across all wetlands. In contrast, soil electrical conductivity (EC) and cation exchange capacity (CEC) were not significantly affected by altitude, although their values varied from 0.29\u0026ndash; 0.37 dS m⁻\u0026sup1; and 20.86\u0026ndash; 31.04 cmol (P⁺) kg⁻\u0026sup1; across wetlands in both seasons. Soil organic carbon (SOC) at the 0\u0026ndash;15 cm depth showed significant variation along the altitudinal gradient, ranging from 68.59\u0026ndash; 95.80 g kg⁻\u0026sup1;. The highest SOC levels were recorded at Koting-Sa-ha Ramosetsana (KSHM) during both the standard wet season (95.03 g kg⁻\u0026sup1;) and the peak wet season (95.80 g kg⁻\u0026sup1;), while Tenesolo (TNL) exhibited the lowest SOC among the wetlands. Overall, SOC tended to increase with elevation in both seasons. Seasonally, EC, CEC, and pH showed a slight decline from the standard wet season to the peak wet season, whereas SOC increased (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSeasonal variation of soil physical and hydraulic properties as influenced by altitudinal gradients\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"10\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" morerows=\"1\" nameend=\"c2\" namest=\"c1\" rowspan=\"2\"\u003e \u003cp\u003eTreatment(s)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c6\" namest=\"c3\"\u003e \u003cp\u003eSpring (Standard wet season)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c10\" namest=\"c7\"\u003e \u003cp\u003eSummer (Peak wet season)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003cp\u003e(Mg m\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eKsat\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eIR\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eMWHC\u003c/p\u003e \u003cp\u003e(%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003cp\u003e(Mg m\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eKsat\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eIR\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eMWHC\u003c/p\u003e \u003cp\u003e(%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWetlands\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAltitude (m) asl\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003ecm hr\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c9\" namest=\"c8\"\u003e \u003cp\u003ecm hr\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhorong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2500\u0026ndash;2550\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e47.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2.60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e50.04\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTenesolo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2552\u0026ndash;2600\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.74\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e41.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e43.90\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhamoqana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2839\u0026ndash;2880\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e43.89\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e46.20\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhalong-La- Lichelete\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2891\u0026ndash;2995\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e44.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e47.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLets'eng- La-Likhama\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3040\u0026ndash;3080\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e49.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e51.82\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKoting-Sa-ha Ramosetsana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3087\u0026ndash;3155\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e54.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e2.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e2.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e57.51\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSE (m)\u0026plusmn;\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.059\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.120\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.094\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.723\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.016\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.111\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.085\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e1.814\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eCD (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.019\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.374\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.292\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.368\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.049\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.345\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.266\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e5.651\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"10\"\u003eCD (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) = Critical Difference at less than 5% probability level; SE(m) = Standard Error of the mean; BD= Bulk density, IR= Infiltration rate, Ksat= Saturated\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003ehydraulic conductivity, MWHC= Maximum water holding capacity, NS\u0026thinsp;=\u0026thinsp;Non-significant.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSeasonal variation of physico-chemical properties of the soil as influenced by altitudinal gradients\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"10\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" morerows=\"1\" nameend=\"c2\" namest=\"c1\" rowspan=\"2\"\u003e \u003cp\u003eTreatment(s)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c6\" namest=\"c3\"\u003e \u003cp\u003eSpring (Standard wet season)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c10\" namest=\"c7\"\u003e \u003cp\u003eSummer (Peak wet season)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eCEC\u003c/p\u003e \u003cp\u003e(cmol) (P\u003csup\u003e+\u003c/sup\u003e) kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eEC\u003c/p\u003e \u003cp\u003e(dS m\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSOC\u003c/p\u003e \u003cp\u003e(g kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eCEC\u003c/p\u003e \u003cp\u003e(cmol) (P\u003csup\u003e+\u003c/sup\u003e) kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eEC\u003c/p\u003e \u003cp\u003e(dS m\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSOC\u003c/p\u003e \u003cp\u003e(g kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWetlands\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAltitude (m) asl\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhorong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2500\u0026ndash;2550\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e28.94\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e83.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e27.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e84.67\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTenesolo\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2552\u0026ndash;2600\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e25.66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e68.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e21.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e69.14\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhamoqana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2839\u0026ndash;2880\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e28.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.93\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e72.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e26.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e73.24\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKhalong-La- Lichelete\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2891\u0026ndash;2995\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e27.93\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e79.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e20.86\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e80.27\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLets'eng- La-Likhama\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3040\u0026ndash;3080\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e26.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e93.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e25.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e94.34\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKoting-Sa-ha Ramosetsana\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3087\u0026ndash;3155\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e31.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e95.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e30.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e6.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e95.80\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSE (m)\u0026plusmn;\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.882\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.074\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.014\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.384\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.939\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.072\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.014\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e8.28\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eCD (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.231\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.197\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.223\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eNS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e18.24\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"10\"\u003eCD (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) = Critical Difference at less than 5% probability level; SE(m) = Standard Error of the mean; asl= Above sea level; CEC= Cation exchange capacity; EC= Electrical\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003econductivity; SOC=Soil organic carbon; NS\u0026thinsp;=\u0026thinsp;Non-significant.\u003c/p\u003e \u003cp\u003e \u003cb\u003e4.4. Pearson\u0026rsquo;s correlation of measured soil attributes during spring (Standard Wet Season) and summer (Peak Wet Season) to evaluate seasonal differences in soil parameter relationships\u003c/b\u003e \u003c/p\u003e \u003cp\u003eDuring the standard wet season, sand particles exhibited a significant negative correlation with both silt (\u0026minus;\u0026thinsp;0.904*) and clay (\u0026minus;\u0026thinsp;0.884*) (Table\u0026nbsp;\u003cspan refid=\"Tab8\" class=\"InternalRef\"\u003e7\u003c/span\u003ea). Clay content showed a significant negative relationship with bulk density (\u0026minus;\u0026thinsp;0.834*), but was positively correlated with infiltration rate (IR) (0.909*) and soil organic carbon (SOC) (0.812*). Bulk density also displayed a significant negative correlation with IR (\u0026minus;\u0026thinsp;0.856*). Maximum water holding capacity (MWHC) demonstrated a highly significant positive association with saturated hydraulic conductivity (Ksat) (0.991**). In addition, IR showed a highly significant positive correlation with SOC (0.993**), whereas electrical conductivity (EC) was significantly and negatively correlated with SOC (\u0026minus;\u0026thinsp;0.845*).\u003c/p\u003e \u003cp\u003eDuring the peak wet season, sand exhibited a significant negative correlation with both silt and clay (Table\u0026nbsp;\u003cspan refid=\"Tab8\" class=\"InternalRef\"\u003e7\u003c/span\u003eb). Clay content showed a significant positive correlation with bulk density (BD) (0.855*) and a significant negative correlation with cation exchange capacity (CEC) (\u0026minus;\u0026thinsp;0.819*). Bulk density was also significantly and negatively correlated with maximum water holding capacity (MWHC) (\u0026minus;\u0026thinsp;0.885*) and CEC (\u0026minus;\u0026thinsp;0.934**). MWHC demonstrated a highly significant positive association with soil organic carbon (SOC) (0.932**). Saturated hydraulic conductivity (Ksat) showed a highly significant positive correlation with infiltration rate (IR) (0.993**) and a significant positive correlation with CEC (0.848*). Electrical conductivity (EC) also exhibited a significant negative correlation with SOC (\u0026minus;\u0026thinsp;0.837*), consistent with the trend observed during the standard wet season.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003e4.5. Seasonal Index of soil physical, hydraulic and physico-chemical attributes\u003c/h2\u003e \u003cp\u003eDuring spring (standard wet season) and summer (peak wet season), several soil properties showed slight deviations from the seasonal average (Figs.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e, \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e and \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Sand content was 0.2% higher in spring and 0.2% lower in summer relative to the seasonal average (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Similarly, silt content was 1% lower in spring but 1% higher in summer. Clay particles were 1.3% above the seasonal average in spring and 1.3% below it in summer. Bulk density was 6.8% higher in spring and correspondingly lower in summer (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Saturated hydraulic conductivity showed a 13% increase in spring and a decrease of the same magnitude in summer. Infiltration rate was 5.1% lower in spring and higher in summer. Maximum water holding capacity was 2.6% lower in spring but higher in summer compared with the seasonal average.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab7\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 7\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003ea: Correlation among the soil physical, hydraulic and physico-chemical properties as influenced by altitudinal gradients\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"12\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e\u003cb\u003eSand\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSand\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSilt\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eClay\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eMWHC\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKsat\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eIR\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003epH\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eEC\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eCEC\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c12\"\u003e \u003cp\u003eSOC\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSilt\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.904\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eClay\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.884\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.600\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eBD\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.627\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.313\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.834\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eMWHC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.073\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.468\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.376\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.642\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eKsat\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.145\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.526\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.305\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.580\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.991\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eIR\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.736\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.430\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.909\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.856\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.554\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.470\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003epH\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.251\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.464\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.038\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.011\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.465\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.556\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.084\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eEC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.719\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.647\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.638\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.491\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.034\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.161\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.712\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.687\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eCEC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.622\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.639\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.467\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.352\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.028\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.036\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.524\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.125\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e-0.311\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSOC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.641\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.355\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.812\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.788\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.495\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.383\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.933\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.294\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e-0.845\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.266\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003ein spring (standard wet season).\u003c/p\u003e \u003cp\u003e*Correlation is significant (p\u0026thinsp;\u0026lt;\u0026thinsp;0.01), **Correlation is highly significant (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05), BD= Bulk density, IR= Infiltration rate, Ksat= Saturated hydraulic conductivity, EC= Electrical conductivity,\u003c/p\u003e \u003cp\u003eCEC= Cation Exchange Capacity and SOC= Soil organic carbon, MWHC= Maximum water holding capacity, NS\u0026thinsp;=\u0026thinsp;Non-significant.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab8\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 7\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eb: Correlation among the soil physical, hydraulic and physico-chemical properties as influenced by altitudinal gradients in summer (peak wet season).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"12\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e 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colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eMWHC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.428\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.099\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.698\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.885\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eKsat\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.400\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.302\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.400\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e 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align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.694\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.741\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.678\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.588\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eCEC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.697\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.426\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.819\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.934\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.782\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.848\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.800\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.039\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e-0.718\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSOC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.274\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.105\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.650\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-0.777\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.932\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.482\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.392\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.516\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e-0.837\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.705\u003csup\u003eNS\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eRefer to Table\u0026nbsp;\u003cspan refid=\"Tab8\" class=\"InternalRef\"\u003e7\u003c/span\u003ea for abbreviations in the footnotes.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSoil pH was 1.5% higher in spring and lower in summer, while electrical conductivity was 2.5% higher in spring and lower in summer (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Cation exchange capacity was 1.5% lower in spring but higher in summer, and soil organic carbon was 0.4% lower in spring and slightly higher in summer relative to the seasonal average.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussions","content":"\u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003e5.1. Seasonal variation of soil particle size distribution, physico-chemical, physical and hydraulic properties across altitudinal gradients\u003c/h2\u003e \u003cp\u003eThe significant variation in soil particle size distribution (PSD) along the altitudinal gradient highlights the important influence of terrain, hydrology, and weathering processes in shaping alpine wetland soils. Soil properties are widely recognized to develop through the interaction of environmental factors such as climate, relief, organisms, parent material, and time, as described in the classical soil-forming factor framework proposed by Hans Jenny. In mountainous environments, elevation modifies climatic variables including temperature and precipitation and also influences vegetation patterns and erosion intensity. These changes affect the mobilization, transport, and deposition of soil particles across landscapes (Liu et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In the present study, sand content decreased with increasing altitude, while silt and clay fractions increased. This pattern suggests that finer particles tend to accumulate in higher elevation wetlands where runoff energy is relatively low and depositional processes dominate. Conversely, lower elevation sites tend to retain coarser particles because stronger runoff selectively removes finer sediments through erosion. Similar altitudinal patterns have been documented in other mountain ecosystems where enhanced chemical weathering and sediment deposition promote the accumulation of finer soil particles at higher elevations (Nthebere et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeasonal variations in PSD further reflect the role of rainfall dynamics in sediment redistribution within wetland environments. The slight increase in sand content and reduction in clay from the standard wet season to the peak wet season may be associated with intensified rainfall and increased runoff during the summer period. Under such conditions, fine particles, particularly clay, are more susceptible to dispersion and transport within saturated soils, whereas sand particles tend to remain relatively stable due to their larger size and mass (Six \u003cem\u003eet al\u003c/em\u003e., 2016; Akor et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The relatively stable silt fraction indicates that seasonal sediment transport processes mainly affected the finest soil particles. Consequently, the predominance of sandy loam textures at lower elevations and loam textures at higher elevations reflects the long-term interaction between erosion, sediment deposition, and hydrological processes along the altitudinal gradient.\u003c/p\u003e \u003cp\u003eThe spatial distribution of soil physical and hydraulic properties also corresponded with variations in elevation and soil texture. Bulk density generally decreased with increasing altitude, whereas maximum water holding capacity (MWHC), saturated hydraulic conductivity (Ksat), and infiltration rate (IR) tended to increase at higher elevation wetlands, particularly at Koting-Sa-ha Ramosetsana. Lower bulk density values at higher elevations are commonly associated with greater organic matter accumulation and increased pore space, which reduce soil compaction and improve soil structure (Lal, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Wetland soils typically accumulate substantial organic residues because prolonged soil saturation slows microbial decomposition. The accumulation of organic matter promotes aggregate stability and the formation of interconnected pore networks, thereby enhancing water retention and hydraulic conductivity (Reddy and DeLaune, 2017). Consequently, the higher MWHC, Ksat, and IR recorded at upper elevation wetlands likely reflect improved soil structure and increased porosity associated with organic-rich soils.\u003c/p\u003e \u003cp\u003eSeasonal differences in these physical and hydraulic parameters further illustrate the influence of soil moisture dynamics. Higher bulk density, infiltration rate, and hydraulic conductivity during the standard wet season may indicate relatively stable soil structure before prolonged saturation occurs. As rainfall intensifies during the peak wet season, extended waterlogging may lead to swelling of fine particles and partial blockage of soil pores, thereby reducing infiltration and hydraulic conductivity (Lal, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In contrast, maximum water holding capacity increased during the peak wet season, reflecting the enhanced moisture retention capacity of soils under saturated conditions. These responses highlight the close interaction between hydrological processes and soil physical functioning in wetland ecosystems.\u003c/p\u003e \u003cp\u003eThe chemical characteristics of the soils also exhibited patterns associated with elevation and seasonal dynamics. Soil pH values remained within the slightly acidic range across the wetlands and varied significantly along the altitudinal gradient. Lower pH values at higher elevations are commonly associated with increased rainfall and enhanced leaching of base cations such as calcium, magnesium, potassium, and sodium (Lal, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Furthermore, the decomposition of organic residues in wetland soils produces organic acids that can contribute to soil acidification. Electrical conductivity (EC) and cation exchange capacity (CEC), however, did not show strong altitudinal responses, suggesting that these properties may be more strongly influenced by soil mineral composition and organic matter content than by elevation alone. The relatively low EC values observed across the wetlands indicate minimal salt accumulation, which is typical of environments characterized by high precipitation and continuous water movement (Reddy and DeLaune, 2017).\u003c/p\u003e \u003cp\u003eSoil organic carbon (SOC) exhibited a clear increase with altitude, with the highest concentrations observed at Koting-Sa-ha Ramosetsana and the lowest at Tenesolo. This trend is consistent with ecological processes commonly observed in mountainous environments where cooler temperatures and higher moisture levels reduce microbial activity and slow organic matter decomposition (Blagodatskaya et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Lal, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Wetland conditions further intensify this effect because saturated soils create anaerobic environments that limit microbial mineralization of organic matter (Reddy and DeLaune, 2017). As a result, organic carbon accumulates more readily over time. Elevated SOC levels at higher elevation wetlands therefore reflect the combined influence of cooler climatic conditions, persistent soil moisture, and slower decomposition rates. In addition to reflecting biogeochemical processes, SOC is widely recognized as an important indicator of wetland ecological health. High SOC concentrations often indicate strong carbon sequestration capacity, sustained vegetation productivity, and stable hydrological conditions (Mitsch et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The relatively high SOC values observed at the upper elevation wetlands therefore suggest that these sites may possess comparatively better ecological condition and greater carbon storage potential.\u003c/p\u003e \u003cp\u003eSeasonal variations in soil chemical properties were less pronounced than the altitudinal differences. Slight decreases in EC, CEC, and pH during the peak wet season may be attributed to increased leaching under conditions of intense rainfall and prolonged soil saturation. Conversely, SOC showed a modest increase during the peak wet season, likely reflecting enhanced plant growth and greater organic residue inputs during the summer growing period combined with slower decomposition under waterlogged conditions (Lal, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAlthough the results revealed clear altitudinal and seasonal patterns, several limitations should be considered. Soil sampling was conducted only during the standard wet season and the peak wet season, and therefore does not capture soil dynamics during the dry season when moisture depletion, oxidation of organic matter, and soil compaction may significantly alter soil characteristics. In addition, variations in water table depth and short-term rainfall intensity, important drivers of wetland soil processes, were not directly measured. Future studies incorporating year-round monitoring and detailed hydrological measurements would provide a more comprehensive understanding of how climatic variability and elevation interact to influence soil processes in alpine wetlands.\u003c/p\u003e \u003cp\u003eOverall, the findings demonstrate that both elevation and seasonal hydrological conditions play critical roles in shaping the physical, hydraulic, and chemical properties of alpine wetland soils. Variations in soil texture, bulk density, water retention capacity, and organic carbon storage reflect the underlying soil-forming processes operating within mountain wetland landscapes and provide valuable insight into the functioning and ecological condition of these sensitive ecosystems.\u003c/p\u003e \u003cp\u003e \u003cb\u003e5.2. Seasonal Index of soil physical, hydraulic and physico-chemical attributes and Pearson\u0026rsquo;s correlation of measured soil attributes during late spring and summer\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThe correlation patterns observed between soil properties during the standard wet season (spring) and the peak wet season (summer) reflect the dynamic hydrological and structural processes that regulate soil functioning in alpine wetlands. The strong negative correlation of sand with both silt and clay in both seasons indicates the inherent textural trade-off within soil particle size distribution, where an increase in coarse particles occurs at the expense of finer fractions. Such relationships are typical in wetland soils and influence water retention, infiltration, and nutrient dynamics along altitudinal gradients (Zhao et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Li \u003cem\u003eet al\u003c/em\u003e., 2020).\u003c/p\u003e \u003cp\u003eDuring the standard wet season, clay showed a significant negative relationship with bulk density but positive correlations with infiltration rate and soil organic carbon. This pattern suggests that finer particles combined with organic matter promote the formation of stable soil aggregates and pore networks, which improve infiltration and reduce soil compaction. In alpine wetlands, higher SOC associated with clay fractions enhances soil structure and water transmission, thereby supporting wetland hydrological regulation and ecosystem functioning (Minasny and McBratney, 2018). The negative correlation between bulk density and infiltration rate further confirms that compacted soils restrict water movement, while less dense soils allow rapid infiltration and groundwater recharge.\u003c/p\u003e \u003cp\u003eThe highly significant association between maximum water holding capacity and saturated hydraulic conductivity suggests that soils with greater pore space can simultaneously retain and transmit water efficiently. This is particularly important in alpine wetlands, where seasonal water storage regulates downstream hydrology and maintains wetland resilience during fluctuating moisture conditions (Hribljan \u003cem\u003eet al\u003c/em\u003e., 2016). The strong positive correlation between infiltration rate and SOC also highlights the ecological role of organic matter in improving soil porosity and permeability. Conversely, the negative correlation between electrical conductivity and SOC suggests that higher organic matter may dilute soluble salts or promote leaching, reducing salinity levels within wetland soils (Yang and Guo, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDuring the peak wet season, several correlations shifted slightly, indicating seasonal hydrological influence. The positive relationship between clay and bulk density suggests temporary compaction of fine particles under prolonged saturation. At the same time, the negative correlation between clay and cation exchange capacity may reflect dilution or redistribution of exchangeable ions under high moisture conditions. The strong negative relationships of bulk density with both maximum water holding capacity and CEC emphasize that compact soils limit water retention and nutrient exchange processes, which are critical for wetland productivity. Hydraulic parameters also showed strong seasonal coupling. The highly significant positive relationship between saturated hydraulic conductivity and infiltration rate indicates synchronized water movement through soil pores during peak rainfall periods. Additionally, the positive association of saturated hydraulic conductivity with CEC suggests that soils capable of transmitting water efficiently may also facilitate nutrient mobility and exchange.\u003c/p\u003e \u003cp\u003eThe slight seasonal deviations in particle size distribution and soil physical properties indicate that alpine wetlands respond sensitively to seasonal hydrological cycles. Lower bulk density and higher infiltration and water holding capacity in summer reflect improved soil structure under sustained moisture conditions. Collectively, these correlations demonstrate that interactions among soil texture, organic carbon, and hydraulic properties regulate water storage, nutrient retention, and ecological stability in alpine wetland systems.\u003c/p\u003e \u003c/div\u003e"},{"header":"Implications and policy recommendation","content":"\u003cp\u003eThe results highlight several implications for the effective conservation and management of alpine wetlands in Lesotho. Variations in soil texture, hydraulic behaviour, and soil organic carbon (SOC) along the altitudinal gradient suggest that wetlands located at higher elevations play a crucial ecological role as natural reservoirs for water storage and as significant carbon sinks. These functions contribute to regulating hydrological processes and maintaining ecological balance in mountainous landscapes. Nevertheless, the presence of fine soil particles that can easily be displaced during intense rainfall or surface disturbance indicates that these wetlands remain highly sensitive to external pressures such as excessive grazing, livestock trampling, and poorly managed land-use activities. Consequently, environmental policies should place greater emphasis on safeguarding high-altitude wetlands where elevated SOC levels and improved soil structure reflect relatively healthier ecosystem conditions and stronger carbon storage capacity. Practical management measures may include regulating grazing intensity, rehabilitating degraded wetland areas, and creating protective buffer zones around vulnerable wetland systems to minimize soil compaction and sediment displacement. Furthermore, incorporating systematic wetland monitoring into national environmental management programmes would enable continuous evaluation of soil quality and hydrological performance. Considering the strong influence of seasonal rainfall variability on soil processes, adaptive land management approaches that respond to climatic fluctuations are equally important. Encouraging community participation in wetland stewardship and promoting sustainable rangeland practices can strengthen the long-term resilience of alpine wetlands while sustaining essential ecosystem services such as water regulation, biodiversity conservation, and carbon retention within Lesotho\u0026rsquo;s highland regions.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIn conclusion, the results indicated that both elevation and seasonal hydrological conditions play a major role in shaping soil texture, hydraulic characteristics, and chemical properties in alpine wetlands. The higher altitude wetland, Koting-Sa-ha Ramosetsana, contained finer soil fractions, lower bulk density, greater water retention capacity, and higher soil organic carbon compared with Tenesolo, suggesting better soil structure, stronger carbon storage potential, and improved ecological status. Seasonal rainfall patterns also influenced sediment movement and soil water dynamics. These outcomes highlight the hydrological and ecological significance of these alpine wetlands. Future studies should incorporate continuous annual monitoring, vegetation assessments, and water-table measurements to improve understanding of long-term soil and wetland processes.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eCRediT authorship contribution statement\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eKnight Nthebere and Mosiuoa Mochala: Writing \u0026ndash; review and editing, Writing \u0026ndash; original draft. Knight Nthebere: Conceptualization. Knight Nthebere and Mosiuoa Mochala: Formal analysis. Knight Nthebere and Mosiuoa Mochala: Methodology and Data curation. Knight Nthebere: Investigation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe research project was funded by the Deutsche Gesellschaft f\u0026uuml;r Internationale Zusammenarbeit (GIZ) and the German Government. The funds were managed by the WaterNet Trust.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData\u0026nbsp;Availability Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eData are provided within the manuscript or Supplementary Information files.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe\u0026nbsp;authors\u0026nbsp;are\u0026nbsp;extremely\u0026nbsp;thankful to\u0026nbsp;the Government of Lesotho, in general, and the ReNOKA movement in particular, which initiated and coordinated the implementation of the Integrated Catchment Management Programme in Lesotho, the EU, Deutsche Gesellschaft f\u0026uuml;r Internationale Zusammenarbeit (GIZ) and the German Government for providing funds for the implementation of this study, as well as WaterNet for managing the research fund.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflicts of Interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAkor S, Flores AN, Rudisill W, Bergstrom A, McNamara JP (2025) Impact of cloud microphysics schemes and boundary conditions on modeled snowpack in mountain watersheds. \u003cem\u003eAuthorea Preprints\u003c/em\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBlagodatskaya Е, Blagodatsky S, Khomyakov N, Myachina O, Kuzyakov Y (2016) Temperature sensitivity and enzymatic mechanisms of soil organic matter decomposition along an altitudinal gradient on Mount Kilimanjaro. 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Water 15(6):1140\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003e\u003cdiv class=\"InlineMediaObject\"\u003e\u003c/div\u003e Supplementary Materials\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSupplementary Fig 1: Overview of Tenesolo (a), Khorong (b), Khamoqana (c), Khalong-La- Lichete (d), Lets\u0026rsquo;eng-La-Likhama (e) and Koting-Sa- ha Ramosetsana (f). Lon= Longitude, Lat= Latitude\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSupplementary Table S. Ecological class utilized for percentage score (PES) evaluations of inland aquatic ecosystems in South Africa, along with the applicable range of PESs for each class (Kleynhans, Macfarlane et al (1996) 2009) [color coding is per the River Eco-Status Monitoring Programme (REMP) of DWS]\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"National University of Lesotho","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Seasonal soil dynamics, seasonal soil index, soil attributes, climate adaptation, ecosystem-based adaptability, high altitudinal gradient","lastPublishedDoi":"10.21203/rs.3.rs-9212062/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9212062/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eSeasonal fluctuations significantly influenced soil physical, hydraulic, and physico-chemical properties in Lesotho\u0026rsquo;s alpine wetlands, yet these ecosystems remain largely understudied. This study evaluated composite seasonal index and examined soil variation and seasonal changes in soil attributes across altitudes (2500\u0026ndash;3155 m a.s.l.), equivalent to wetlands from three sub-catchments (blocks): Khorong and Tenesolo (Senqunyane), Khamoqana and Khalong-la-Lichelete (Sani), and Lets\u0026rsquo;eng-la-Likhama and Koting-Sa-ha Ramosetsana (Khubelu), during the standard (spring, September 2024) and peak (summer, February 2025) wet seasons. The soil samples were collected in September 2024 (standard) and February 2025 (peak) wet seasons and analyzed for bulk density (BD), saturated hydraulic conductivity (Ksat), infiltration rate (IR), water holding capacity (WHC), texture, pH, electrical conductivity (EC), cation exchange capacity (CEC) and soil organic carbon (SOC) following standard procedures. Soil texture exhibited clear altitudinal trends: sand decreased from 64.97% at lower elevations to 39.79% upslope, whereas silt and clay increased, resulting in sandy-loam at lower and loam at higher sites. Seasonal variations, though subtle, were measurable: sand (\u0026plusmn;\u0026thinsp;0.2%), silt (\u0026plusmn;\u0026thinsp;1%), clay (\u0026plusmn;\u0026thinsp;1.3%), BD (6.8%), Ksat (13%), IR (5.1%), and WHC (2.6%). SOC rose slightly (~\u0026thinsp;0.4%) in summer, peaking at Koting-Sa-ha Ramosetsana. Correlation analysis indicated strong negative relationships between sand and both clay and silt, while BD inversely related to IR. Positive associations were observed between WHC and Ksat, and IR correlated closely with SOC. Overall, these results underscore that seasonal hydrological dynamics and elevation jointly shape soil structure, water movement, and carbon storage in Lesotho\u0026rsquo;s alpine-wetlands, emphasizing their ecological sensitivity and need for targeted conservation.\u003c/p\u003e","manuscriptTitle":"Seasonal Dynamics of Physical, Hydraulic, and Physico-Chemical Attributes of the Soil across Altitudinal Gradients in the Alpine Wetlands of Lesotho","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-25 08:56:49","doi":"10.21203/rs.3.rs-9212062/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"9c623b2c-92c4-40ea-94c8-a982152a3315","owner":[],"postedDate":"March 25th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":65053523,"name":"Agroecology"}],"tags":[],"updatedAt":"2026-03-25T08:56:49+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-25 08:56:49","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9212062","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9212062","identity":"rs-9212062","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}