# Genesis and Classification of Termite-Mediated Soils along Toposequences in a Semiarid Area of Southeast Ethiopia

Authors: Abinet Bekele; Sheleme Beyene; Alemayehu Kiflu; Fantaw Yimer
Journal: Applied and Environmental Soil Science
 (2023)

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## Abstract

Despite the ecosystem functioning they provide, termite pedoturbation along toposequence is overlooked in the genesis of semiarid soils. Therefore, we aimed to describe morphological and physicochemical properties that lead to the classification of termite-mediated soils. In this study, representative pedons, one on each slope class, were described and classified for five different topographical positions, and the soil properties of genetic horizons were compared to those obtained from respective mounds. The result showed that the soils were heavily manipulated by termites except for the pedon at the toe slope. Cambisols were formed on the summit and back slope and resulted from slow pedogenic processes. Luvisols on the toe slope showed redoximorphic features, and gleization and clay synthesis formed the soil, while the upward movement of coarse particles enhanced textural differentiation. Luvisols at the foot of the slope are formed by the partial destruction of iron-bearing minerals accompanied by eluviation-illuviation processes. Accumulation of calcium carbonate following calcification formed Calcisols on the bottom slope. Comparing the mounds and reference pedons, much of the mound’s soil is mined from the subsoil, usually from B horizons. However, their influence on soil properties depended mainly on the topography. Moreover, the morphological and physicochemical properties of the studied pedons exhibited various degrees of variation along topography and clearly showed topographic effects. In conclusion, termites can be a potent mediator of soil genesis across toposequences, and their activities should be considered in the classification and management of semiarid soils. A further retrospective examination of micromorphological evidence is recommended to support this finding.


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## Body

## 1. Introduction

Mound-forming termites and large epigeal mounds are the common features of the landscape in most semiarid regions [1, 2]. Termites through their pedoturbation drastically change soil properties and influence soil genesis [3, 4]. The mechanism by which termites affect soil formation is by altering the type of soil that had formed years ago, e.g., from Alisols and Acrisols to umbric properties [4], by extracting clay-enriched materials from the subsoil, e.g., from the Kandic horizon [5] and the Argic horizon [4], by promoting soil homogenization [3], by influencing soil formation processes [6], and by influencing morphological properties [7]. Another important impact is the tunneling and refilling of the subsoil with topsoil [8] and by modifying the mineralogical composition of the soil [9].Topography is also important in explaining soil variability and pathways of soil transformation. According to Breuning-Madsen et al. [10], macrofauna-induced continuous surface exposure on sloppy landscapes together with episodic precipitation can cause significant downward soil transport. Thuyne and Verrecchia [11] showed that the termite-induced sediment accumulation rate reached up to 1 mm·ha−1·y−1. In such a landscape, profile development can be much more complex than vertically up or down processes since mixing by soil fauna interacts with lateral processes [12].Although the role of termites in soil formation has been recognized for a long time, only a few studies have been conducted on the pedogenesis of termite-affected soils [3, 4, 13, 14]. In most of these studies, the diversity of pedogenetic processes involved in the genesis of termite-mediated soils is neglected. In Ethiopia, where mound-forming termites predominantly inhibit arid to semiarid ecosystems, termite-mediated soils have been characterized for a specific location [15], and there are still limited studies on the soil genesis of termite-affected soils. Regardless of the differences in soils across topography and the tendency of termites to selectively mine fine particles at different depths [16], the extent to which epigeal termite mounds resemble the reference soils across toposequence is largely unknown. On the other hand, the study of soil characterization and classification that focuses on moisture-deficient areas is limited, and to our knowledge, only the study conducted by Ahmed et al. [17] had initiated in the past.Since soil is central in an ecosystem, having insight into termite-mediated soils makes this study uniquely contribute to the knowledge of semiarid soils. Additionally, understanding soil-forming processes could also contribute to and have a genetic signal for better soil characterization [18]. Therefore, this study was initiated to characterize the morphological and physicochemical properties of termite-mediated soils, quantify the properties of termite mounds, and compare them with those obtained from genetic horizons which were not influenced by termites.


## 2. Materials and Methods

### 2.1. Description of the Study Area

Epigeal termite mounds are the common features of the landscape in a semiarid area of southeast Ethiopia, starting fromDugda Dawa in the north along the highway to Moyale in the south. Geographically, this area lies between 3°53′ to 5°22′ N and 38°16′ to 38°33′ E. Two districts, Dugda Dawa in the north and Miyo in the south, were deliberately selected for this study because the epigeal mounds in the two areas have different physical characteristics. Both study areas are characterized by bimodal rainfall, with the main rainy season occurring between March and May and the short rainy season occurring between October and November. The mean annual rainfall of the Dugda Dawa district is about 711 mm with a mean annual temperature of 19.9°C ranging from 13.3°C to 26.6°C. The rainfall in the Miyo area is 345.5 mm, and the mean annual temperature is 23.9°C, ranging from 18.4°C to 29.5°C. The geology of the parent material is dominated by quartzo-feldspathic gneisses, gneiss and granulite, magnetite-quartz-feldspar gneisses, quaternary metamorphic rock, and sedimentary deposits [19]. The geographic location and the geological conditions of the study area are shown in Figure 1.Figure 1
The study area. (a) The districts in Oromia Regional State, Ethiopia. (b, c) The map of the study area with geological features of theDugda Dawa and Miyo districts, respectively. Geological information is derived from Ó Dochartaigh [19].


### 2.2. Soil Pedon Description and Sampling

Representative toposequences with widespread termite mounds were selected at theIlala-Sara and Silala sites in Dugda Dawa and Miyo districts, respectively. The toposequence in Ilala-Sara was grouped into the summit, back slope, and toe slope, while the toposequence in Silala included the foot slope and bottom slope. A pedon was excavated at each topographical position and described in situ following the Guidelines for Soil Description [20], and soil color was determined based on Munsell Soil Color Charts [21]. The soils were classified according to [18]. The location of each pedon was georeferenced using a portable GPS (62-GARMIN).Additionally, abundance and other termite mound attributes were estimated per topographical position, and termite specimens were collected for taxonomic identification. At each position, three active representative mounds were selected at a distance of 10 m from the reference pedon and considered for soil sampling. A composite soil sample was collected per mound after the entire epigeal mound had been broken up and thoroughly mixed, and a total of three composites were made for termite mounds within a topographical position. However, soil samples were not collected from mounds at toe slopes, as these mounds were less abundant and inactive. All composite soil samples from genetic horizons and mounds were separately tagged and transported to the laboratory.


### 2.3. Soil Sample Preparation and Analyses

All analyses were performed for the fine Earth fractions (<2 mm). Soil particle size was analyzed by the Bouyoucos hydrometer method [22]. Soil pH and electrical conductivity (EC) were measured in a 1 : 2.5 soil-to-water suspension. Total organic carbon (OC) was determined by wet oxidation in acidified dichromate [23], while available phosphorus was determined using the Olsen method [24]. Exchangeable cations (K+, Na+, Ca2+, and Mg2+) and cation exchangeable capacity (CEC) were extracted using 1 M NH4OAc at pH 7.0 [25]. While Ca2+ and Mg2+ in the leachate were measured with an atomic absorption spectrophotometer (AAS), K+ and Na+ were determined with a flame photometer. Calcium carbonate was estimated by HCl neutralization and titration with NaOH [26] and reported as calcium carbonate equivalent (CCE). Available micronutrients (Zn, Mn, Fe, and Cu) were extracted using the diethylenetriaminepentaacetic acid method, and the contents in the leachate were measured using AAS.


## 2.1. Description of the Study Area

Epigeal termite mounds are the common features of the landscape in a semiarid area of southeast Ethiopia, starting fromDugda Dawa in the north along the highway to Moyale in the south. Geographically, this area lies between 3°53′ to 5°22′ N and 38°16′ to 38°33′ E. Two districts, Dugda Dawa in the north and Miyo in the south, were deliberately selected for this study because the epigeal mounds in the two areas have different physical characteristics. Both study areas are characterized by bimodal rainfall, with the main rainy season occurring between March and May and the short rainy season occurring between October and November. The mean annual rainfall of the Dugda Dawa district is about 711 mm with a mean annual temperature of 19.9°C ranging from 13.3°C to 26.6°C. The rainfall in the Miyo area is 345.5 mm, and the mean annual temperature is 23.9°C, ranging from 18.4°C to 29.5°C. The geology of the parent material is dominated by quartzo-feldspathic gneisses, gneiss and granulite, magnetite-quartz-feldspar gneisses, quaternary metamorphic rock, and sedimentary deposits [19]. The geographic location and the geological conditions of the study area are shown in Figure 1.Figure 1
The study area. (a) The districts in Oromia Regional State, Ethiopia. (b, c) The map of the study area with geological features of theDugda Dawa and Miyo districts, respectively. Geological information is derived from Ó Dochartaigh [19].


## 2.2. Soil Pedon Description and Sampling

Representative toposequences with widespread termite mounds were selected at theIlala-Sara and Silala sites in Dugda Dawa and Miyo districts, respectively. The toposequence in Ilala-Sara was grouped into the summit, back slope, and toe slope, while the toposequence in Silala included the foot slope and bottom slope. A pedon was excavated at each topographical position and described in situ following the Guidelines for Soil Description [20], and soil color was determined based on Munsell Soil Color Charts [21]. The soils were classified according to [18]. The location of each pedon was georeferenced using a portable GPS (62-GARMIN).Additionally, abundance and other termite mound attributes were estimated per topographical position, and termite specimens were collected for taxonomic identification. At each position, three active representative mounds were selected at a distance of 10 m from the reference pedon and considered for soil sampling. A composite soil sample was collected per mound after the entire epigeal mound had been broken up and thoroughly mixed, and a total of three composites were made for termite mounds within a topographical position. However, soil samples were not collected from mounds at toe slopes, as these mounds were less abundant and inactive. All composite soil samples from genetic horizons and mounds were separately tagged and transported to the laboratory.


## 2.3. Soil Sample Preparation and Analyses

All analyses were performed for the fine Earth fractions (<2 mm). Soil particle size was analyzed by the Bouyoucos hydrometer method [22]. Soil pH and electrical conductivity (EC) were measured in a 1 : 2.5 soil-to-water suspension. Total organic carbon (OC) was determined by wet oxidation in acidified dichromate [23], while available phosphorus was determined using the Olsen method [24]. Exchangeable cations (K+, Na+, Ca2+, and Mg2+) and cation exchangeable capacity (CEC) were extracted using 1 M NH4OAc at pH 7.0 [25]. While Ca2+ and Mg2+ in the leachate were measured with an atomic absorption spectrophotometer (AAS), K+ and Na+ were determined with a flame photometer. Calcium carbonate was estimated by HCl neutralization and titration with NaOH [26] and reported as calcium carbonate equivalent (CCE). Available micronutrients (Zn, Mn, Fe, and Cu) were extracted using the diethylenetriaminepentaacetic acid method, and the contents in the leachate were measured using AAS.


## 3. Results and Discussion

### 3.1. Sites and Morphological Characteristics of Termite Mounds

The site characteristics of the pedons showed variations in drainage, degree of erosion, and surface stoniness (Table1). Pedons D01, D02, and D03 are located on the summit, back slope, and toe slope positions, respectively, while pedon M01 is located on the foot slope and pedon M02 is found on the bottom slope. Pedons D01 and D02 were formed from saprolitic materials, while pedon D03 was developed from alluvium deposits set back from slope sediments. Pedon M01 was derived from finely textured eluvium residuum, while pedon M02 was established from lacustrine deposits. Although termite mounds inhibited the survey sites, pedon M01 indicated intense termite activity, as sheetings were formed overnight in an open pit.Table 1
Site characterization of the soil profiles across toposequence in Ilala-Sara and Silala, southeast Ethiopia.
ParametersIlala-Sara, Dugda DawaSilala, MiyoPedon,D01Pedon,D02Pedon,D03Pedon,M01Pedon,M02Coordinatea5°23′0.41″N, 38°16′19.85″E5°22′52.33″N, 38°16′13.26″E5°22′46.41″N, 38°16′8.42″E3° 53′ 41.81″N, 38° 33' 54.81″E3° 53′ 24.27″N, 38° 33′ 47.64″EElevationb16311592159013421333Major landformPlainPlainAlluvial plainPlainPlainLand surfaceGently undulatingGently undulatingNearly levelVery gently slopingLevelled land surfaceTopographic positionSummitBack slopeToe slopeFoot slopeBottomSlope gradient (%)450.71.60.2Slope formsStraight convexStraight convexStraight concaveStraightStraightAspectSoutheastSoutheastSoutheastSoutheastSoutheastDrainage classWell-drainedWell-drainedSomewhat poorly drainedWell-drainedWell-drainedEvidence of erosion/depositionSlight evidence of erosion damaged 25 to 50% of the total areaSlight evidence of erosion damaged more than 50% of the total area with some evidence of gulliesSlight evidence of deposition by water covering about 5–10% of the total areaSlight evidence of erosion damaged more than 50% of the total areaNo evidence of erosionRock outcrops/stoninessCommon stonesMany coarse gravelNoneNoneVery few bouldersNatural vegetationSemideciduous woodlandSemideciduous woodlandSemideciduous woodlandSemideciduous woodlandHerbaceous short grasslandLand-useOpen grazing landOpen grazing landOpen grazing landOpen grazing landOpen grazing landGeologyCenozoic-quaternary-Holocene quartzo-feldspathic gneisses, gneiss and granuliteCenozoic-quaternary-Holocene quartzo-feldspathic gneisses, gneiss, and granulitePrecambrian lower Paleozoic magnetite-quartz-feldspar gneiss: Quartz, feldspar, and magnetiteLower Paleozoic quaternary metamorphic rockPrecambrian Jurassic sedimentary rockParent materialsSaproliteSaproliteFine-textured materials derived from alluvium depositUnconsolidated, weathered, and eluvium, finely textured residuumClay and silt fractions predominate lacustrine deposits of base-rich materialsaUTM coordinate, Zone 37°N, datum WGS 84. bmeter above sea level.Two termite genera representing the studied toposequences were identified. The genusMacrotermes built open, cathedral-shaped mounds that reached up to 4 m high at all slope positions, except at the bottom slope, where it formed closed conical mounds with a mean height of up to 3.2 m (Table 2). The other termite genus, Odontotermes, built closed, dome-shaped mounds at the toe slope with a mean height of up to 2.2 m. Both genera are fungus-growing termites that are commonly reported in Africa [27] as well as the study area [1, 28]. These genera influence savanna soils much more than the other termite groups [11]. The abundance of Macrotermes mounds reached up to 23 mounds ha−1, while the Odontotermes genera had about 4 mounds ha−1. Macrotermes deposit up to 12.51 m3·ha−1, demonstrating an important role for termites in soil pedogenesis. According to Wilkinson et al. [29], abundant and massive mounds on the summit and back slopes may be related to saprolite because saprolites buffer water availability during climate fluctuations for soil organisms [30]. In general, mounds on gentle slopes had a larger structure than mounds on a nearly flat landscape. Larger mounds have stable soil chemistry [31]. Accordingly, termite pedoturbation is intense in the gently undulating landscapes of the study area. In support of this result, Abe et al. [6] reported that the influence of termites on pedogenesis is more evident in inclined than level positions.Table 2
The characteristics of termite mounds vary across toposequence in Ilala-Sara and Silala, southeast Ethiopia.
Topographic positionsMound density (ha−1)Mound vitalityMound height (m)Mean basal radius of mound (m)Volume of mound (m3 mound−1)Mass of the selected mounds (t mound−1)ActiveDeadIlala-Sara, Dugda Dawa districtSummit141133.621.7010.9514.67Back slope181354.671.6012.5116.77Toe slope4142.20.962.122.42Silala, Miyo districtFoot slope171253.31.063.885.20Bottom slope231763.20.882.592.96


### 3.2. Differentiation of Soil Morphology across the Toposequences

The soils of the study area were characterized by deep pedons, typically over 200 cm (Table3). The thinnest A horizon was observed in pedon D02, while the thickest surface horizon was in pedon M01. Pedons with deeper depths may be due to the nature of the parent materials, which favor permeability to the depth at which termites search for fine particles. Thick surface horizons could be acquired due to erosion off the mounds and galleries which redistributed over the landscape and resulted in deeper soil depth [32]. In contrast, thin horizons could be related to the removal of surface materials as a result of topographic effects.Table 3
Morphological soil properties of the pedons across toposequences in Ilala-Sara and Silala, southeast Ethiopia.
HorizonDepth (cm)Horizon boundaryaMunsell color notationSoil structural units’ grade, size, and typebConsistencecRoots’ abundance and sizedRock fragment abundance, classification, and weatheringeMoistDryDryMoistStickinessPlasticityPedon D01, Ilala-Sara, ∼ gently undulating summitA0–12DW2.5 YR4/42.5 YR4/8WE, ME, CRSOVFRSSTNPLM and MC—AB12–30CW2.5 YR3/62.5 YR4/8WE, CO, GRSOVFRSSTSPLV and FVF, MG, and FBw30–73CW2.5 YR4/42.5 YR4/6ST, FC, RSEHAEFISSTPLV and VFM, CGS, and FBC73–140GW7.5 YR4/67.5 R5/6WE, ME, SBSHAEFISSTSPLV and VFFe, MCG, and WCr140–200+—10 R4/410 R4/3WE, VF, MASOLONSTNPLV and VFFe, MCG, and WPedon D02, Ilala-Sara site, ∼ gently undulating back slopeA0–7GW2.5 YR4/42.5YR5/4WE, VF, CRSOLONSTNPLC and VFVF, FG, and FAB7–95CW2.5YR3/62.5YR4/8ST, CO, RSHAVFINSTSPLF and VFA, CGS, and FBw95–140CW2.5YR4/82.5YR5/8WE, FI, ABSHAFRSSTSPLF and VFFe, FMG, and FBC140–170GW2.5YR4/82.5YR5/8WE, FM, ABSHAVFRNSTSPLV and VFVF, FG, and WCr170–200+—2.5YR5/82.5YR6/8WE, FI, MASOFRNSTNPL——Pedon D03, Ilala-Sara site, ∼ nearly levelled toe slopeA0–10CW2.5YR4/42.5YR5/4WE, ME, SBSOLOSSTNPLC, VF—AB10–40CW2.5YR3/62.5YR4/8WE, FM, SBSOLONSTNPLF, FVF, MG, FBg40–70AS7.5YR4/17.5YR5/3WE, ME, PRSHAVFRSSTSPLF, M—Bt70–200+—7.5YR3/17.5YR4/1MO, VC, ABHAFINSTVPLV, F—Pedon M01, Silala site, ∼ very gentle foot slopeA0–10CS2.5YR4/42.5YR4/6MO, ME, GRSHAVFRSSTNPLM, FM—AB10–55GS10R3/610R4/6WE, FM, SBSHAVFRSSTSPLC, FM—Bt55–90GS2.5YR5/42.5YR3/6WE, FM, ABSHAVFRSSTSPLF, F—Bo90–200+—2.5YR3/610R3/6WE, CO, PR ⟶ ABHAFRSSTSPL——Pedon M02, Silala site, ∼ levelled land surfaceA0–18CS5YR4/27.5YR6/1WE, FM, SBSOVFRSSTPLM, FF/MC—AB18–42CS2.5YR5/110YR5/2WE, FM, SBSOLOSSTSPLC, FMC, CG, FBk42–70CS5Y6/15Y8/1WE, MC, SBSOVFRSSTSPLC, FMVF, FG, FCk70–120GS2.5Y8/25Y8/1ST, CO, MAHAFINSTNPLV, VFD, MG, WCkm120–200+—2.5Y7/15Y8/1ST, CO, MAHAFINSTNPL—D, CG, FaAS: abrupt smooth, CS: clear smooth, CW: clear wavy, DW: diffuse wavy, GS: gradual smooth, and GW: gradual wavy. bAB: angular blocky, CO: coarse/thick, FC: fine to coarse, FI: fine/thin, FM: fine and medium, GR: granular, MA: massive, ME: medium, MO: moderate, MVC: medium to very coarse, PR: prismatic, RS: rock structure, SB: subangular blocky, ST: strong, VC: very coarse/thick, VF: very fine/thin, and WE: weak. cEFI: extremely firm, EHA: extremely hard, FI: firm, FR: friable, HA: hard, LO: loose, NPL: nonplastic, NST: nonsticky, PL: plastic, SHA: slightly hard, SO: soft, SPL: slightly plastic, SST: slightly sticky, VFI: very firm, VFR: very friable, VPL: very plastic, and VST: very sticky. dC: common/coarse, F: fine/few, FF: few fine, FM: fine to medium, M: many/medium, MC: medium to coarse, V: very, and VF: very fine. eA: abundant, C: common, CG: coarse gravel, CGS: coarse gravel and stones, D: dominant, F: fresh or slightly weathered, Fe: few, FG: fine gravel, FMG: fine and medium gravel, M: many, MCG: medium and coarse gravel, MG: medium gravel, VF: very few, and W: weathered.Horizontal boundaries varied along the toposequences, and pedonsD01 and D02 had diffuse and gradual wavy boundaries at the surface and gradual wavy boundaries at the subsurface (Table 3). While gradual wavy boundaries may be associated with underlying saprolitic materials, wavy horizon topography likely accentuates the presence of lamellae [33]. According to FAO [34], biological pedoturbation leads to diffuse or gradual horizon boundaries. A pedon D03 had a clear wavy horizon boundary superimposed on an abrupt smooth horizon, reflecting the mode of deposition [35]. Pedons M01 and M02 had clear to gradual smooth horizon boundaries, which were probably facilitated by termite pedoturbation along with fine to medium grassroots [36, 37]. In general, gradual distinctness in most pedons is attributed to the systematic and continuous supply of soil materials [38], whereas wavy and smooth boundaries signify the removal of soil materials due to topography and reworked soil horizons [39].Pedons vary greatly in color, and soil redness decreases down the slope (Table3). Consistent with the findings by Jimoh et al. [40], small amounts of OC mixed with soil particles resulted in a reddish-brown surface soil color across the toposequences that becomes lighter with decreasing OC in the subsurface of the pedons. In most pedons, the hue pattern began to emerge by 2.5YR, although there were some irregularities. The consistent hue pattern could be due to a termite-induced translocation of soil materials. All examined pedons had a low value and high chroma, except for pedon M02, which tended to show a high value and low chroma. Conspicuously, low value and high chroma without apparent iron illuviation in the pedons D01, D02, and M01 may be related to illuvial sesquioxide accumulation. Due to the proximity of pedon D03 to seasonal floods, the reddish-brown color of the surface horizon might be owing to oxidized iron, while a very dark gray subsurface horizon indicates iron reduction. This result is consistent with the findings of Romanens et al. [41] who observed Fe coatings that give a reddish color to the top horizons without any contact with underlying horizons. Regarding the soil color of termite mounds, representative mounds were red (2.5 YR 4/6, dry) on the summit and back slope, dark red (2.5 YR 3/6, dry) on the foot slope, and white (5 YR 8/1, dry) on the bottom slope. The similarity of termite mound color to subsurface horizons in reference pedons supports the fact that termites collect subsurface soil particles for mound formation [3, 7, 15].The surface soil of the studied pedons had a weak fine to medium granular, crumbly, and subangular blocky texture that changed to a weak medium weak fine to very coarse angular blocky texture, a weak medium subangular blocky or prismatic texture, and very strong and very coarse rock in the B horizons and massive rocks in the C horizons (Table3). The structural unit of the topsoil may be related to termite pedoturbation. This is consistent with the recent findings of De Freitas et al. [3], which indicated that soil that has been worked a lot by termites exhibited granular and small subangular block structures. According to Bera et al. [7], subangular structures with a reddish color predominated also in the subsoil, as pedoturbation decreased with depth. In accordance with Dinssa and Elias [42], the angular block structure formed in the subsoil of D02 could be due to an overlying dense soil layer, while that of D03 and M01 might be related to enriched clay particles, reduced OM, and root distribution.The soil consistency was less differentiated among the topographical positions (Table3). With some exceptions, all pedons are dominated by soft, slightly hard to hard dry consistencies, with moist consistencies ranging from loose, very friable to extremely firm. These pedons were generally marked by a slight stickiness ranging from nonplastic to plastic. The granular or fine subangular blocky structures probably favored the friable and very friable consistencies at the surface horizons of the pedons D01, M01, and M02 due to moderate to high OC levels. The representation of wet consistence in the subsoil of most horizons is attributed to reduced OM and clay accumulations [42]. A soft, dry consistency in the surface horizon of M02 may underpin secondary carbonate. The lack of stickiness and plasticity in some horizons could be related to the amount of clay particles in the soil. In contrast, that of pedon M02 could be attributed to indurate carbonate.The surface layers of pedonsD01 and D02 had very few fine gravels (>2 mm), while gravelly horizons were exposed to the surface in pedon D02 (Table 3). The subsurface horizons of these pedons were marked by abundant coarse gravels and stones that changed to very few fine gravels with depth. Pedon D03 had very few medium gravels near the surface layer that decreased to zero with depth. In pedon M02, rock fragments were distinguished in most of the subsurface horizons, and these exceeded more than 80% in the lowest horizons. Gravelly horizons in pedons D01 and D02 are probably due to fragments caused by dominant physical weathering and are not related to termite stone lines. This is because a termite-related stone layer is identified by the distinguished chemical properties of the soil overlying the layer [14]. In these pedons, a reduced fragment distribution with depth indicates downslope transported soil materials [33]. The fragments are exposed to the surface layers on the back slope indicating the influence of topography since surface erosion is higher than the rate of replenishment.


### 3.3. Physical Properties of Termite-Mediated Soils

Both soil particle sizes (≤2 mm) and silt: clay ratios are irregularly distributed over topographical positions (Table4). The absence of a definite pattern of soil particle sizes is probably attributed to surface soil erosion and subsequent accumulation which may influence pedogenic processes [43]. Sand particles were gradually decreased in all pedons, except for pedon M02 which exhibited an increasing pattern with depth. In line with these results, a gradually decreasing sand pattern was also reported by Bonifacio et al. [44]. According to Jimoh et al. [40], the trends of sand distribution in pedon M02 might be attributed to irregular deposition. Based on the rating by Hazelton and Murphy [45], all pedons had a high to very high sand content, while the silt content was classified as very low to low. The high to very high sand content could be related to the influence of parent materials. Given that the soils are in the moisture deficit region, such low silt contents were common. According to Daher et al. [46], negligible ornithogenesis and sulphides probably impede the advanced weathering of silts and clays in semiarid regions.Table 4
Selected soil physicochemical properties of the termite mounds (TM,n = 3, ± standard deviation) and reference pedons across toposequences in Ilala-Sara and Silala, southeast Ethiopia.
HorizonsDepth (cm)Sand (>63μm)Silt (2–63μm)Clay (<2μm)Texture classSilt : clay ratiopH (1 : 2.5 H2O)CCE (g kg−1)EC (mS cm−1)OC (g kg−1)Avail. P (mg kg−1)CEC (cmolc kg−1)g kg−1 of fine EarthPedon D01, Ilala-Sara, ∼ gently undulating summitA0–1268060260Sandy clay loam0.235.741.4912.484.815.20AB12–3050080420Sandy clay0.195.951.8313.264.2610.00Bw30–73440120440Clay0.275.491.2911.709.1710.60BC73–140510130360Sandy clay0.365.921.348.7808.629.00Cr140–200+720140140Sandy loam1.0a06.061.996.6305.576.00TM—480 ± 1.15140 ± 2.00380 ± 2.31Sandy clay0.376.83 ± 0.3113.21 ± 0.0222.88 ± 0.249.13 ± 0.9632.67 ± 2.19Pedon D02, Ilala-Sara site, ∼ gently undulating back slopeA0–778080140Sandy loam0.575.690.9211.707.925.20AB7–9558080340Sandy clay loam0.245.210.8011.706.77.60Bw95–14036020620Clay0.036.031.089.756.544.80BC140–17048060460Sandy clay0.135.40.682.346.79.20Cr170–200+540100360Sandy clay0.285.10.623.906.48.60TM—527 ± 1.33180 ± 2.00293 ± 1.76Sandy clay loam0.616.28 ± 0.1346.14 ± 0.1720.41 ± 0.4011.97 ± 1.1128.67 ± 0.88Pedon D03, Ilala-Sara site, ∼ nearly levelled toe slopeA0–1074040220Sandy clay loam0.185.821.2012.486.387.80AB10–4074040220Sandy clay loam0.185.730.8710.539.176.20Bg40–70660100240Sandy clay loam0.425.630.6011.7012.367.20Bt70–200+340140520Clay0.275.951.279.3610.6926.40Pedon M01, Silala site, ∼ very gentle foot slopeA0–10580200220Sandy clay loam0.916.682.1116.385.029.20AB10–55520100380Sandy clay0.266.010.8012.093.229.80Bt55–90380140480Clay0.296.420.827.802.814.20Bo90–200+480260260Sandy clay loam1.007.222.347.804.3917.40TM—500 ± 1.16167 ± 1.76333 ± 2.67Sandy clay loam0.506.17 ± 0.1214.09 ± 0.0815.73 ± 0.1913.19 ± 0.9029 ± 3.22Pedon M02, Silala site, ∼ levelled land surfaceA0–18480260260Sandy clay loam1.007.60265.003.6623.405.7814.20AB18–42560220220Sandy clay loam1.007.39525.003.7619.506.8910.00Bk42–70480240280Sandy clay loam0.867.62685.003.7713.658.759.40Ck70–120550150300Sandy clay loam0.507.79680.003.9714.439.638.20Ckm120–200+74040220Sandy clay loam0.188.20565.006.4113.654.7416.40TM—353 ± 1.76200 ± 0.00447 ± 1.76Clay0.457.28 ± 0.31663.33 ± 3.758.73 ± 0.0421.58 ± 0.4517.1 ± 1.0528.33 ± 1.76A steady increase in clay content was evident in the pedonsD01, D02, and M01, showing accumulation with depth. The peak clay content of pedon M01 could be related to clay illuviation, while that of pedons D01 and D02 might have been inherited from the saprolite materials, which also resulted in a low silt-to-clay ratio (Table 4). This could be attributed to the fact that saprolites are more clay-rich because the particles are trapped before seeping into the deeper layer [47]. Clay particles varied rapidly with depth in the pedon D03, and such a wetting-front pattern could be associated with heavy illuviation [48]. Based on the CEC of clay, however, the absence of 2 : 1 clay in the upslope soil (pedons D01 and D02) implies that clay accumulation in pedon D03 is not of colluvial origin, whereas the irregular clay pattern within this pedon would not support in situ clay formation [49]. Based on these views, it appears that clay synthesis in the pedon D03 comes from alluvial deposits. On the other hand, clay content showed minimum and maximum peaks with depth in the pedon M02. According to Minasny et al. [48], soil attributes can show a minimal depth pattern when surface mixing is excessive and a translocation process is operating. In this pedon, consistent texture throughout the depth might be inherited from carbonate-rich parent materials [41].The average clay content of the mounds at the summit, back slope, and foot slope was reduced by 14%, 112%, and 44% compared to the maximum content in pedonsD01, D02, and M01, respectively. While the clay content of the mounds was 50% higher than the maximum clay content in the subsurface of pedon M02. In this study, termite pedoturbation affected soil particle-sizes by enriching mounds with silt and clay compared to reference pedons, but the magnitude was related to the topographical positions. Consistent with this study, Freitas et al. [3] reported that a wide range of particles ranging from silt to clay could be selected for mound building. Clay enrichment in termite mounds is related to translocated mica and smectite or saprolitic materials from the subsoil [50]. According to Jouquet et al. [5], high clay content in termite mounds could arise from the dissolution of plant matter, while resilication is also another process, particularly for smectite.Except at the bottom slope, the studied mounds were predominantly sandy clay loam in texture, and termite mound soils were not different from matched reference pedons, corroborating the findings of Freitas et al. [3]. According to Abe and Wakatsuki [49], the clay mineralogy of the termite mound soil is also very similar to adjacent bulk soils. A contrasting mound soil texture, as with the pedon D02, implies that termites mine and translocate soil particles depending on the nature of the subsoil. In addition, the silt-to-clay ratio of termite mounds ranged from 0.37 to 0.61 which was comparable to the ratio obtained from reference pedons (Table 4). This supports previous studies [50, 51] and indicates that termites relocate fine particles from the subsurface of the respective reference soils across toposequences.


### 3.4. Chemical Properties of Termite-Mediated Soils

Soil pH (1 : 2.5 H2O) varied across the toposequences (Table 4), which could be attributed to the leaching of cations from the upslope and the subsequent accumulation at the nearly level positions [52]. Moderately acidic pH increased with depth in pedon D01 and became strongly acidic at the subsurface of pedon D02, while it was uniformly distributed in pedon D03. The soil pH was neutral at the surface, slightly increased with depth in pedon M01, and changed to moderately alkaline in pedon M02. Apart from pedon M02, the acidic soil surface could be related to the leaching of basic cations, while limited leaching due to herbaceous vegetation probably maintained a near neutral soil reaction in pedon M01, whereas the alkalinity of M02 could be attributed to the nature of the parent material. In general, the comparable soil pH within the pedons likely indicates slow weathering and soil development. The soil pH values of termite mounds ranged from slightly acidic to neutral which were not similar to the respective pedons D01 and D02 but similar to the subsurface horizons of the pedons M01 and M02 (Table 4). The soil pH of mounds depends on the corresponding pH values of the reference soil at a given topographical position. This is consistent with the findings of Sarcinelli et al. [32]. According to Bera et al. [7], the pH of termite mounds is attributed to cation-clay accumulation and the rate of OM decomposition in the mounds.While no carbonate was present in all other soils, the soils in the pedonM02 invariably exhibited strong effervescence with dilute HCl (10%) and showed calcium carbonate that rapidly increased with depth (Table 4). Termite mounds at the same position had CCE that was 2.5 times higher than the respective surface soil and comparable to the subsurface of the reference pedon. The accumulation of carbonate within a pedon could be inherited from parent materials and other associated rocks. The CCE-depth relationship also supports the premise that it is geological rather than pedogenic since weathering of the bedrock results in substantial Ca2+ [41]. According to Buss et al. [53], the carbonate incidence near the surface soil suggests a limited percolation of water to leach carbonate. The distinct features of CCE in mounds indicate that termite pedoturbation concentrates carbonate and relays it to bulk soils. This result is consistent with the observation of Mcauliffe et al. [54] who reported the enrichment of mounds with carbonate.Soil EC content showed an irregular pattern across topographical positions owing to leaching and accumulation of basic cations (Table4). Apart from pedon M02, the EC in the horizons of the pedons was below 2.34 mS cm−1. For the pedon M02, the EC value of 8.66 mS cm−1 in the surface soil decreased gradually with depth. In general, negligible EC differences in the soils of most pedons indicate limited eluviation and illuviation processes, while the amounts in the pedon M02 could be related to the substantial basic cations in the soils of the pedon. Compared to the reference pedon, the average EC of mounds was almost seven times higher than the maximum value for soils in the pedons D01 and M01 and 43 times higher in the pedon D02 while comparable in the pedon M02. By and large, the higher EC of termite mounds compared to bulk soils could be attributed to more soluble ions in the biogenic structure [55].The soil OC varied consistently along the toposequences (Table4). This corroborates with the findings of Sarcinelli et al. [32] which suggest that similar compositions of termite foraging material result in comparable OC content across topographical positions. Except for the pedon D03 which had a wetting-front pattern, the OC gradually decreased with depth. The OC of mounds at the summit and back slope exceeded the higher content of the surface horizons by more than 70%. However, the OC content of the mounds was lower than that of the surface bulk soils of the pedons M01 and M02. While wetting-front OC trends might be attributed to slow translocation, the gradual decreasing pattern probably indicates the accumulation of OC through grass decay. Based on the OC-to-clay index stated by Merante et al. [56], the high to moderate OC content in pedon M02 could also be associated with a complex OC with clay content, which could lead to C stabilization. Regarding the OC levels of the mounds, the result is consistent with the observation of Obi et al. [57] and demonstrates topographic effects. In addition to the high mineralization rate in a semiarid area, the fact that fungus-growing termites mineralize about 20% of the total OC [11] probably results in a low OC content.The available P was less than 10.69 mg·kg−1 across the studied pedons and irregularly distributed across toposequences (Table 4). The distribution pattern of P showed accumulation with depth in the pedons D01, D03, and M02, while its value gradually decreased in the pedons D02 and M01. Compared to the reference pedon, the average P content of the mounds was higher by 51%, 163%, and 78% than the maximum values of the soil horizons in the pedons D02, M01, and M02, respectively. In general, low available P could be attributed to insoluble Ca phosphates comprising the calcareous materials in the pedon M02, while the low levels in the other pedons might be due to sorption by sesquioxides. According to Dinssa and Elias [42], a decreasing P-trend with depth could be associated with reduced OM and P fixation. The highly available P of termite mounds could be related to the incorporation of OM into mounds [4]. However, Bera et al. [7] reported fewer contents in mounds arguing that P content depends on the termite feeding habit and the soil materials. The P content of the mound also depends on the amount of clay, pH, types, and termite species involved in mound formations [11]. In our case, the viable explanation for the highly available P content of mounds is probably related to the soil pH nearing neutral reaction, clay content, and long-term accumulation of OM leading to humification which in turn favors the release of P.The CEC of the soils varied irregularly across the toposequences (Table4). In pedons D01 and D02, CEC had an irregular pattern with depth. With a rapidly changing pattern, the pedons D03 and M01 had increasing CEC values with depth, while pedon D03 had substantially high CEC in the lowest horizon. The CEC gradually decreased with depth and accumulated in the lowest horizon of the pedon M02. A low CEC in the surface soil could be associated with a low OM content, while a moderate to high CEC in the subsurface could result from clay accumulation. A distinctly high CEC in the pedon D03 could be due to clay synthesis (r > 0.99∗∗, not shown here) and the accumulation of Ca2+ and Mg2+ (r = 0.99∗∗). On the other hand, the considerable CEC in the pedon M02 could be associated with substantial clay and Ca2+ in the soils [41]. The CEC values of the mounds on the summit and back slope were three times, while that of the mounds at the foot slope and bottom slope were two-fold higher than the maximum CEC in the respective horizons of the reference pedons. The higher CEC of termite mounds might be independent of OM (r = 0.19 ns), and it is rather related to the presence of 2 : 1 clay. This is because the CEC of both soil and clay is higher than what would be expected for soils dominated by kaolinite, which suggests the presence of montmorillonite clay in the soil of termite mounds and related to the subsoil of respective pedons. This corroborates with the findings of Abe and Wakatsuki [49] who observed minor modification of clay mineralogy in Macrotermes mounds compared to their respective soils. Consequently, this clay probably explains the high CEC of termite mounds.While Ca2+, Mg2+, and K+ showed an increasing trend, Na+ exhibited decreasing patterns across topographical positions (Table 5). The increasing pattern of Ca2+ and Mg2+ down the slope could be due to lateral movement. Exchangeable Ca2+ was less than 10 cmolc kg−1 in all pedons except for pedon M02 and a distinctly high value in the lowermost horizon of pedon D03. Both Ca2+ and Mg2+ had an irregular pattern with depth in the pedons D01 and D02 and M02, rapidly changed with depth in the pedon D03, and showed accumulation in the subsurface of the pedon M01. The high Ca2+ in the pedon D03 could be related to the CaCO3 accumulation [53] and that of the pedon M02 could be attributed to the weathering of carbonate-enriched parent materials [41]. Exchangeable K+ gradually increased in the pedons D01 and D02, while the other pedons gradually decreased with depth. In all pedons, Na+ had an irregular decreasing trend with depth except within pedon D03 which had some enrichment. The increase of K+ with depth in the pedons D01 and D02 could be related to the leaching of cations [40]. Relatively, the high K+ in the pedon D03 is probably due to the accumulation of mica at that position, while the Na+ accumulation in the lowest horizon of this pedon may be due to its greater mobility. The Ca > Mg > K > Na order in the studied pedons might be due to the release of these ions by weathering before the rocks form. The major cations that dominated the exchange complex of the pedons are consistent with the findings by Beyene [58].Table 5
Exchangeable cations and available micronutrients of the termite mounds (TM,n = 3, ± standard deviation) and reference profiles across toposequences in Ilala-Sara and Silala, southeast Ethiopia.
HorizonsSampling depth (cm)Exchangeable cationsAvailable micronutrientsCa2+Mg2+K+Na+FeMnZnCucmolc kg−1mg kg−1Profile D01, Ilala-Sara, ∼ gently undulating summitA0–120.470.400.190.147.0810.300.940.50AB12–300.910.770.190.1210.539.271.390.71Bw30–730.650.480.160.135.775.821.390.64BC73–1401.370.460.260.116.515.391.220.57Cr140–200+1.010.750.270.1210.951.750.140.36TM6.61 ± 0.640.47 ± 0.120.13 ± 0.030.13 ± 0.04————Profile D02, Ilala-Sara site, ∼ gently undulating back slopeA0–71.680.510.110.1511.275.360.270.38AB7–951.210.610.150.159.466.730.090.31Bw95–1401.420.460.120.162.891.020.790.33BC140–1701.240.690.160.1311.331.820.420.36Cr170–200+1.310.530.190.149.442.020.220.36TM19.15 ± 1.310.72 ± 0.140.13 ± 0.020.10 ± 0.00————Profile D03, Ilala-Sara site, ∼ nearly levelled toe slopeA0–101.781.080.380.1013.659.950.520.40AB10–401.671.130.210.0618.2015.980.530.45Bg40–701.190.870.290.0725.385.470.230.54Bt70–200+12.223.200.220.1920.081.940.630.97Profile M01, Silala site, ∼ very gentle foot slopeA0–103.930.330.180.1532.861.932.070.72AB10–552.360.250.150.1242.609.261.741.23Bt55–904.351.300.170.1737.286.691.581.13Bo90–200+5.531.720.150.1431.283.431.240.78TM8.69 ± 0.791.17 ± 0.270.23 ± 0.020.14 ± 0.04————Profile M02, Silala site, ∼ levelled bottom slopeA0–1851.141.500.150.1330.705.191.870.50AB18–4251.470.700.170.1720.262.751.320.42Bk42–7054.651.710.150.1229.325.111.720.43Ck70–12047.124.480.150.1021.304.490.290.46Ckm120–200+31.464.750.100.0927.246.111.380.41TM70.24 ± 2.153.38 ± 0.120.23 ± 0.030.07 ± 0.03————The mounds at the summit and back slope had 4 to 10 times higher Ca2+, while the mounds at the bottom slope had 57 times higher Ca2+ than the maximum value in the respective reference pedon (Table 5). On the other hand, the Mg2+ contents of the mounds were lower compared to that of the surface soils but comparable to the levels in the subsurface horizon of the reference pedons. The K+ content of mounds decreased up to 52% compared to the maximum values in the soils of the reference pedons D01 and D02, while the K+ content of the mounds was higher by more than 28% in the pedons M01 and M02. The exchangeable Na+ of the mounds was decreased by 7% to 59% compared to the maximum values in the reference pedons. While termite pedoturbation did not affect the Na+ and Mg2+ across the toposequences, their effects were evident for Ca2+ at all locations and K+ at the foot slope and bottom slope positions. In support of this study, Obi et al. [57] also considered Ca2+, Mg2+, and K+ as indicators of pedoturbation.


### 3.5. Available Micronutrients

While Zn and Cu decreased down the slope, Fe and Mn showed irregular patterns across toposequences (Table5). Consistent with Hailu et al. [43], the patterns of Fe and Mn were similar at any given topographical position because of their similar chemical behavior. The micronutrient contents were in the order of Fe > Mn > Zn > Cu. According to Zhu et al. [59], the dominance of Fe at any given topographical position might be due to the dissolution of metal oxide in the clay particles which in turn strongly depends on the slope position while the availability of Zn and Cu related to desorption from clay, carbonate, and OM surface. Available Fe, Mn, and Zn contents showed an irregular pattern with depth in the pedons D01, D02, and M02, probably owing to termite pedoturbation while accumulating with depth in pedon D03. While Fe and Mn accumulated just below the surface layer, Zn gradually decreased with depth in the pedon M01. Available Cu showed a gradual decrease with depth in the pedons D01, D02, and M02, while it gradually increased in the pedons D03 and M01.


### 3.6. Pedogenesis and Classification of the Soils

The subsurface horizon of the pedonsD01 and D02 showed an incipient pedogenetic alteration, with half of the original rock structure being obliterated. Additionally, the horizon thickness and texture of the soils qualify them for a cambic horizon. The cambic horizons were also specifically identified by redder hues (2.5 years) and higher chroma (>6) on the surface than the underlying horizons. On the other hand, sufficiently intense reduction and segregation of iron which resulted in low chroma were indicative of the identification of the cambic horizon in the pedon D03. Prominent clay accumulation in the subsoil horizons of the pedon D03 qualified argic diagnostic horizon. The pedon M01 was differentiated by a redder hue, higher chroma, and an accumulation of clay and sesquioxides. The moist color changes from reddish-brown in the upper to dark reddish-brown at the subsurface horizon substantiating clay illuviation for identification of the argic horizon. Termite pedoturbation [13], shrubs, perennials, and grasses probably resulted in clay accumulation in this pedon. A polyhedral structure in this pedon did not meet the diagnostic features of a nitic horizon. The subsurface horizon of the pedon M02 had more than 15% CCE in the fine Earth fraction which meets the diagnostic criteria for a calcic horizon. According to Macedo et al. [60], the absence of secondary carbonate accumulation may be due to termite pedoturbation. Based on the above diagnostic horizons and using established criteria from IUSS Working Group WRB [18], the pedons D01 and D02 were classified as Cambisols. Argic horizons accompanied by CECs of clay greater than 24 cmolc kg−1 in the pedons D03 and M01 differentiated the soils into Luvisols, whereas the pedon M02 was identified as Calcisols. The principal and supplementary qualifiers are summarized in Table 6.Table 6
Descriptions of diagnostic horizons and soil classification based on the WRB system.
Ilala-Sara, Dugda DawaSilala, MiyoPedon,D01Pedon,D02Pedon,D03Pedon,M01Pedon,M02Diagnostic horizonCambic horizonCambic horizonArgic horizonArgic horizonCalcic horizonIUSS working group WRB [18]Hyperdystric endoskeletic cambisols (clayic and humic)Dystric skeletic cambisols (loamic and humic)Gleyic luvisols (clayic, neocambic, differentic, epidystric, and humic)Chromic luvisols (clayic, epidystric, and humic)Skeletic cambic calcisols (loamic and humic)


### 3.7. Pedogenic Processes

The soils in the surface layers of all pedons showed the feature of termite pedoturbation as traced by granular or subangular blocky structures and gradual or diffuse horizon boundaries except for soils in pedonD03. In addition, incipient pedogenic processes, as shown by little alteration and very few fine root distributions, might be some of the soil-forming processes in the pedons D01 and D02. As stated by Borden et al. [52], the loss of cations through downslope translocation also probably promotes desilication in these two pedons. The evidence of gleization was observed in the pedon D03, but cyclic drying and wetting in the surface and the remains of moisture in the subsoil favored hydrolysis which subsequently derived clay formation. In this pedon, the absence of mottling suggests minimal current pedogenetic activities other than gleying and illuviation processes. According to Bonifacio et al. [44], gleyic properties indicate strong weathering of the iron-bearing minerals probably as evidenced by the depletion of available Fe on the surface of the pedon. In the pedon M01, desilication, braunification, rubification, ferrugation, leucinization, and illuviation might be the most important pedogenic processes. Calcification was the dominant process in the pedon M02 while humification dispersed to a certain extent throughout the solum. Following Deressa et al. [36], there was a textural differentiation in examined pedons except for the pedon M02 which showed a clay contrast index of 0.87 indicating a lower textural gradient. In addition to termite pedoturbation, calcification processes result in homogenous clay distribution within the pedon M02.


## 3.1. Sites and Morphological Characteristics of Termite Mounds

The site characteristics of the pedons showed variations in drainage, degree of erosion, and surface stoniness (Table1). Pedons D01, D02, and D03 are located on the summit, back slope, and toe slope positions, respectively, while pedon M01 is located on the foot slope and pedon M02 is found on the bottom slope. Pedons D01 and D02 were formed from saprolitic materials, while pedon D03 was developed from alluvium deposits set back from slope sediments. Pedon M01 was derived from finely textured eluvium residuum, while pedon M02 was established from lacustrine deposits. Although termite mounds inhibited the survey sites, pedon M01 indicated intense termite activity, as sheetings were formed overnight in an open pit.Table 1
Site characterization of the soil profiles across toposequence in Ilala-Sara and Silala, southeast Ethiopia.
ParametersIlala-Sara, Dugda DawaSilala, MiyoPedon,D01Pedon,D02Pedon,D03Pedon,M01Pedon,M02Coordinatea5°23′0.41″N, 38°16′19.85″E5°22′52.33″N, 38°16′13.26″E5°22′46.41″N, 38°16′8.42″E3° 53′ 41.81″N, 38° 33' 54.81″E3° 53′ 24.27″N, 38° 33′ 47.64″EElevationb16311592159013421333Major landformPlainPlainAlluvial plainPlainPlainLand surfaceGently undulatingGently undulatingNearly levelVery gently slopingLevelled land surfaceTopographic positionSummitBack slopeToe slopeFoot slopeBottomSlope gradient (%)450.71.60.2Slope formsStraight convexStraight convexStraight concaveStraightStraightAspectSoutheastSoutheastSoutheastSoutheastSoutheastDrainage classWell-drainedWell-drainedSomewhat poorly drainedWell-drainedWell-drainedEvidence of erosion/depositionSlight evidence of erosion damaged 25 to 50% of the total areaSlight evidence of erosion damaged more than 50% of the total area with some evidence of gulliesSlight evidence of deposition by water covering about 5–10% of the total areaSlight evidence of erosion damaged more than 50% of the total areaNo evidence of erosionRock outcrops/stoninessCommon stonesMany coarse gravelNoneNoneVery few bouldersNatural vegetationSemideciduous woodlandSemideciduous woodlandSemideciduous woodlandSemideciduous woodlandHerbaceous short grasslandLand-useOpen grazing landOpen grazing landOpen grazing landOpen grazing landOpen grazing landGeologyCenozoic-quaternary-Holocene quartzo-feldspathic gneisses, gneiss and granuliteCenozoic-quaternary-Holocene quartzo-feldspathic gneisses, gneiss, and granulitePrecambrian lower Paleozoic magnetite-quartz-feldspar gneiss: Quartz, feldspar, and magnetiteLower Paleozoic quaternary metamorphic rockPrecambrian Jurassic sedimentary rockParent materialsSaproliteSaproliteFine-textured materials derived from alluvium depositUnconsolidated, weathered, and eluvium, finely textured residuumClay and silt fractions predominate lacustrine deposits of base-rich materialsaUTM coordinate, Zone 37°N, datum WGS 84. bmeter above sea level.Two termite genera representing the studied toposequences were identified. The genusMacrotermes built open, cathedral-shaped mounds that reached up to 4 m high at all slope positions, except at the bottom slope, where it formed closed conical mounds with a mean height of up to 3.2 m (Table 2). The other termite genus, Odontotermes, built closed, dome-shaped mounds at the toe slope with a mean height of up to 2.2 m. Both genera are fungus-growing termites that are commonly reported in Africa [27] as well as the study area [1, 28]. These genera influence savanna soils much more than the other termite groups [11]. The abundance of Macrotermes mounds reached up to 23 mounds ha−1, while the Odontotermes genera had about 4 mounds ha−1. Macrotermes deposit up to 12.51 m3·ha−1, demonstrating an important role for termites in soil pedogenesis. According to Wilkinson et al. [29], abundant and massive mounds on the summit and back slopes may be related to saprolite because saprolites buffer water availability during climate fluctuations for soil organisms [30]. In general, mounds on gentle slopes had a larger structure than mounds on a nearly flat landscape. Larger mounds have stable soil chemistry [31]. Accordingly, termite pedoturbation is intense in the gently undulating landscapes of the study area. In support of this result, Abe et al. [6] reported that the influence of termites on pedogenesis is more evident in inclined than level positions.Table 2
The characteristics of termite mounds vary across toposequence in Ilala-Sara and Silala, southeast Ethiopia.
Topographic positionsMound density (ha−1)Mound vitalityMound height (m)Mean basal radius of mound (m)Volume of mound (m3 mound−1)Mass of the selected mounds (t mound−1)ActiveDeadIlala-Sara, Dugda Dawa districtSummit141133.621.7010.9514.67Back slope181354.671.6012.5116.77Toe slope4142.20.962.122.42Silala, Miyo districtFoot slope171253.31.063.885.20Bottom slope231763.20.882.592.96


## 3.2. Differentiation of Soil Morphology across the Toposequences

The soils of the study area were characterized by deep pedons, typically over 200 cm (Table3). The thinnest A horizon was observed in pedon D02, while the thickest surface horizon was in pedon M01. Pedons with deeper depths may be due to the nature of the parent materials, which favor permeability to the depth at which termites search for fine particles. Thick surface horizons could be acquired due to erosion off the mounds and galleries which redistributed over the landscape and resulted in deeper soil depth [32]. In contrast, thin horizons could be related to the removal of surface materials as a result of topographic effects.Table 3
Morphological soil properties of the pedons across toposequences in Ilala-Sara and Silala, southeast Ethiopia.
HorizonDepth (cm)Horizon boundaryaMunsell color notationSoil structural units’ grade, size, and typebConsistencecRoots’ abundance and sizedRock fragment abundance, classification, and weatheringeMoistDryDryMoistStickinessPlasticityPedon D01, Ilala-Sara, ∼ gently undulating summitA0–12DW2.5 YR4/42.5 YR4/8WE, ME, CRSOVFRSSTNPLM and MC—AB12–30CW2.5 YR3/62.5 YR4/8WE, CO, GRSOVFRSSTSPLV and FVF, MG, and FBw30–73CW2.5 YR4/42.5 YR4/6ST, FC, RSEHAEFISSTPLV and VFM, CGS, and FBC73–140GW7.5 YR4/67.5 R5/6WE, ME, SBSHAEFISSTSPLV and VFFe, MCG, and WCr140–200+—10 R4/410 R4/3WE, VF, MASOLONSTNPLV and VFFe, MCG, and WPedon D02, Ilala-Sara site, ∼ gently undulating back slopeA0–7GW2.5 YR4/42.5YR5/4WE, VF, CRSOLONSTNPLC and VFVF, FG, and FAB7–95CW2.5YR3/62.5YR4/8ST, CO, RSHAVFINSTSPLF and VFA, CGS, and FBw95–140CW2.5YR4/82.5YR5/8WE, FI, ABSHAFRSSTSPLF and VFFe, FMG, and FBC140–170GW2.5YR4/82.5YR5/8WE, FM, ABSHAVFRNSTSPLV and VFVF, FG, and WCr170–200+—2.5YR5/82.5YR6/8WE, FI, MASOFRNSTNPL——Pedon D03, Ilala-Sara site, ∼ nearly levelled toe slopeA0–10CW2.5YR4/42.5YR5/4WE, ME, SBSOLOSSTNPLC, VF—AB10–40CW2.5YR3/62.5YR4/8WE, FM, SBSOLONSTNPLF, FVF, MG, FBg40–70AS7.5YR4/17.5YR5/3WE, ME, PRSHAVFRSSTSPLF, M—Bt70–200+—7.5YR3/17.5YR4/1MO, VC, ABHAFINSTVPLV, F—Pedon M01, Silala site, ∼ very gentle foot slopeA0–10CS2.5YR4/42.5YR4/6MO, ME, GRSHAVFRSSTNPLM, FM—AB10–55GS10R3/610R4/6WE, FM, SBSHAVFRSSTSPLC, FM—Bt55–90GS2.5YR5/42.5YR3/6WE, FM, ABSHAVFRSSTSPLF, F—Bo90–200+—2.5YR3/610R3/6WE, CO, PR ⟶ ABHAFRSSTSPL——Pedon M02, Silala site, ∼ levelled land surfaceA0–18CS5YR4/27.5YR6/1WE, FM, SBSOVFRSSTPLM, FF/MC—AB18–42CS2.5YR5/110YR5/2WE, FM, SBSOLOSSTSPLC, FMC, CG, FBk42–70CS5Y6/15Y8/1WE, MC, SBSOVFRSSTSPLC, FMVF, FG, FCk70–120GS2.5Y8/25Y8/1ST, CO, MAHAFINSTNPLV, VFD, MG, WCkm120–200+—2.5Y7/15Y8/1ST, CO, MAHAFINSTNPL—D, CG, FaAS: abrupt smooth, CS: clear smooth, CW: clear wavy, DW: diffuse wavy, GS: gradual smooth, and GW: gradual wavy. bAB: angular blocky, CO: coarse/thick, FC: fine to coarse, FI: fine/thin, FM: fine and medium, GR: granular, MA: massive, ME: medium, MO: moderate, MVC: medium to very coarse, PR: prismatic, RS: rock structure, SB: subangular blocky, ST: strong, VC: very coarse/thick, VF: very fine/thin, and WE: weak. cEFI: extremely firm, EHA: extremely hard, FI: firm, FR: friable, HA: hard, LO: loose, NPL: nonplastic, NST: nonsticky, PL: plastic, SHA: slightly hard, SO: soft, SPL: slightly plastic, SST: slightly sticky, VFI: very firm, VFR: very friable, VPL: very plastic, and VST: very sticky. dC: common/coarse, F: fine/few, FF: few fine, FM: fine to medium, M: many/medium, MC: medium to coarse, V: very, and VF: very fine. eA: abundant, C: common, CG: coarse gravel, CGS: coarse gravel and stones, D: dominant, F: fresh or slightly weathered, Fe: few, FG: fine gravel, FMG: fine and medium gravel, M: many, MCG: medium and coarse gravel, MG: medium gravel, VF: very few, and W: weathered.Horizontal boundaries varied along the toposequences, and pedonsD01 and D02 had diffuse and gradual wavy boundaries at the surface and gradual wavy boundaries at the subsurface (Table 3). While gradual wavy boundaries may be associated with underlying saprolitic materials, wavy horizon topography likely accentuates the presence of lamellae [33]. According to FAO [34], biological pedoturbation leads to diffuse or gradual horizon boundaries. A pedon D03 had a clear wavy horizon boundary superimposed on an abrupt smooth horizon, reflecting the mode of deposition [35]. Pedons M01 and M02 had clear to gradual smooth horizon boundaries, which were probably facilitated by termite pedoturbation along with fine to medium grassroots [36, 37]. In general, gradual distinctness in most pedons is attributed to the systematic and continuous supply of soil materials [38], whereas wavy and smooth boundaries signify the removal of soil materials due to topography and reworked soil horizons [39].Pedons vary greatly in color, and soil redness decreases down the slope (Table3). Consistent with the findings by Jimoh et al. [40], small amounts of OC mixed with soil particles resulted in a reddish-brown surface soil color across the toposequences that becomes lighter with decreasing OC in the subsurface of the pedons. In most pedons, the hue pattern began to emerge by 2.5YR, although there were some irregularities. The consistent hue pattern could be due to a termite-induced translocation of soil materials. All examined pedons had a low value and high chroma, except for pedon M02, which tended to show a high value and low chroma. Conspicuously, low value and high chroma without apparent iron illuviation in the pedons D01, D02, and M01 may be related to illuvial sesquioxide accumulation. Due to the proximity of pedon D03 to seasonal floods, the reddish-brown color of the surface horizon might be owing to oxidized iron, while a very dark gray subsurface horizon indicates iron reduction. This result is consistent with the findings of Romanens et al. [41] who observed Fe coatings that give a reddish color to the top horizons without any contact with underlying horizons. Regarding the soil color of termite mounds, representative mounds were red (2.5 YR 4/6, dry) on the summit and back slope, dark red (2.5 YR 3/6, dry) on the foot slope, and white (5 YR 8/1, dry) on the bottom slope. The similarity of termite mound color to subsurface horizons in reference pedons supports the fact that termites collect subsurface soil particles for mound formation [3, 7, 15].The surface soil of the studied pedons had a weak fine to medium granular, crumbly, and subangular blocky texture that changed to a weak medium weak fine to very coarse angular blocky texture, a weak medium subangular blocky or prismatic texture, and very strong and very coarse rock in the B horizons and massive rocks in the C horizons (Table3). The structural unit of the topsoil may be related to termite pedoturbation. This is consistent with the recent findings of De Freitas et al. [3], which indicated that soil that has been worked a lot by termites exhibited granular and small subangular block structures. According to Bera et al. [7], subangular structures with a reddish color predominated also in the subsoil, as pedoturbation decreased with depth. In accordance with Dinssa and Elias [42], the angular block structure formed in the subsoil of D02 could be due to an overlying dense soil layer, while that of D03 and M01 might be related to enriched clay particles, reduced OM, and root distribution.The soil consistency was less differentiated among the topographical positions (Table3). With some exceptions, all pedons are dominated by soft, slightly hard to hard dry consistencies, with moist consistencies ranging from loose, very friable to extremely firm. These pedons were generally marked by a slight stickiness ranging from nonplastic to plastic. The granular or fine subangular blocky structures probably favored the friable and very friable consistencies at the surface horizons of the pedons D01, M01, and M02 due to moderate to high OC levels. The representation of wet consistence in the subsoil of most horizons is attributed to reduced OM and clay accumulations [42]. A soft, dry consistency in the surface horizon of M02 may underpin secondary carbonate. The lack of stickiness and plasticity in some horizons could be related to the amount of clay particles in the soil. In contrast, that of pedon M02 could be attributed to indurate carbonate.The surface layers of pedonsD01 and D02 had very few fine gravels (>2 mm), while gravelly horizons were exposed to the surface in pedon D02 (Table 3). The subsurface horizons of these pedons were marked by abundant coarse gravels and stones that changed to very few fine gravels with depth. Pedon D03 had very few medium gravels near the surface layer that decreased to zero with depth. In pedon M02, rock fragments were distinguished in most of the subsurface horizons, and these exceeded more than 80% in the lowest horizons. Gravelly horizons in pedons D01 and D02 are probably due to fragments caused by dominant physical weathering and are not related to termite stone lines. This is because a termite-related stone layer is identified by the distinguished chemical properties of the soil overlying the layer [14]. In these pedons, a reduced fragment distribution with depth indicates downslope transported soil materials [33]. The fragments are exposed to the surface layers on the back slope indicating the influence of topography since surface erosion is higher than the rate of replenishment.


## 3.3. Physical Properties of Termite-Mediated Soils

Both soil particle sizes (≤2 mm) and silt: clay ratios are irregularly distributed over topographical positions (Table4). The absence of a definite pattern of soil particle sizes is probably attributed to surface soil erosion and subsequent accumulation which may influence pedogenic processes [43]. Sand particles were gradually decreased in all pedons, except for pedon M02 which exhibited an increasing pattern with depth. In line with these results, a gradually decreasing sand pattern was also reported by Bonifacio et al. [44]. According to Jimoh et al. [40], the trends of sand distribution in pedon M02 might be attributed to irregular deposition. Based on the rating by Hazelton and Murphy [45], all pedons had a high to very high sand content, while the silt content was classified as very low to low. The high to very high sand content could be related to the influence of parent materials. Given that the soils are in the moisture deficit region, such low silt contents were common. According to Daher et al. [46], negligible ornithogenesis and sulphides probably impede the advanced weathering of silts and clays in semiarid regions.Table 4
Selected soil physicochemical properties of the termite mounds (TM,n = 3, ± standard deviation) and reference pedons across toposequences in Ilala-Sara and Silala, southeast Ethiopia.
HorizonsDepth (cm)Sand (>63μm)Silt (2–63μm)Clay (<2μm)Texture classSilt : clay ratiopH (1 : 2.5 H2O)CCE (g kg−1)EC (mS cm−1)OC (g kg−1)Avail. P (mg kg−1)CEC (cmolc kg−1)g kg−1 of fine EarthPedon D01, Ilala-Sara, ∼ gently undulating summitA0–1268060260Sandy clay loam0.235.741.4912.484.815.20AB12–3050080420Sandy clay0.195.951.8313.264.2610.00Bw30–73440120440Clay0.275.491.2911.709.1710.60BC73–140510130360Sandy clay0.365.921.348.7808.629.00Cr140–200+720140140Sandy loam1.0a06.061.996.6305.576.00TM—480 ± 1.15140 ± 2.00380 ± 2.31Sandy clay0.376.83 ± 0.3113.21 ± 0.0222.88 ± 0.249.13 ± 0.9632.67 ± 2.19Pedon D02, Ilala-Sara site, ∼ gently undulating back slopeA0–778080140Sandy loam0.575.690.9211.707.925.20AB7–9558080340Sandy clay loam0.245.210.8011.706.77.60Bw95–14036020620Clay0.036.031.089.756.544.80BC140–17048060460Sandy clay0.135.40.682.346.79.20Cr170–200+540100360Sandy clay0.285.10.623.906.48.60TM—527 ± 1.33180 ± 2.00293 ± 1.76Sandy clay loam0.616.28 ± 0.1346.14 ± 0.1720.41 ± 0.4011.97 ± 1.1128.67 ± 0.88Pedon D03, Ilala-Sara site, ∼ nearly levelled toe slopeA0–1074040220Sandy clay loam0.185.821.2012.486.387.80AB10–4074040220Sandy clay loam0.185.730.8710.539.176.20Bg40–70660100240Sandy clay loam0.425.630.6011.7012.367.20Bt70–200+340140520Clay0.275.951.279.3610.6926.40Pedon M01, Silala site, ∼ very gentle foot slopeA0–10580200220Sandy clay loam0.916.682.1116.385.029.20AB10–55520100380Sandy clay0.266.010.8012.093.229.80Bt55–90380140480Clay0.296.420.827.802.814.20Bo90–200+480260260Sandy clay loam1.007.222.347.804.3917.40TM—500 ± 1.16167 ± 1.76333 ± 2.67Sandy clay loam0.506.17 ± 0.1214.09 ± 0.0815.73 ± 0.1913.19 ± 0.9029 ± 3.22Pedon M02, Silala site, ∼ levelled land surfaceA0–18480260260Sandy clay loam1.007.60265.003.6623.405.7814.20AB18–42560220220Sandy clay loam1.007.39525.003.7619.506.8910.00Bk42–70480240280Sandy clay loam0.867.62685.003.7713.658.759.40Ck70–120550150300Sandy clay loam0.507.79680.003.9714.439.638.20Ckm120–200+74040220Sandy clay loam0.188.20565.006.4113.654.7416.40TM—353 ± 1.76200 ± 0.00447 ± 1.76Clay0.457.28 ± 0.31663.33 ± 3.758.73 ± 0.0421.58 ± 0.4517.1 ± 1.0528.33 ± 1.76A steady increase in clay content was evident in the pedonsD01, D02, and M01, showing accumulation with depth. The peak clay content of pedon M01 could be related to clay illuviation, while that of pedons D01 and D02 might have been inherited from the saprolite materials, which also resulted in a low silt-to-clay ratio (Table 4). This could be attributed to the fact that saprolites are more clay-rich because the particles are trapped before seeping into the deeper layer [47]. Clay particles varied rapidly with depth in the pedon D03, and such a wetting-front pattern could be associated with heavy illuviation [48]. Based on the CEC of clay, however, the absence of 2 : 1 clay in the upslope soil (pedons D01 and D02) implies that clay accumulation in pedon D03 is not of colluvial origin, whereas the irregular clay pattern within this pedon would not support in situ clay formation [49]. Based on these views, it appears that clay synthesis in the pedon D03 comes from alluvial deposits. On the other hand, clay content showed minimum and maximum peaks with depth in the pedon M02. According to Minasny et al. [48], soil attributes can show a minimal depth pattern when surface mixing is excessive and a translocation process is operating. In this pedon, consistent texture throughout the depth might be inherited from carbonate-rich parent materials [41].The average clay content of the mounds at the summit, back slope, and foot slope was reduced by 14%, 112%, and 44% compared to the maximum content in pedonsD01, D02, and M01, respectively. While the clay content of the mounds was 50% higher than the maximum clay content in the subsurface of pedon M02. In this study, termite pedoturbation affected soil particle-sizes by enriching mounds with silt and clay compared to reference pedons, but the magnitude was related to the topographical positions. Consistent with this study, Freitas et al. [3] reported that a wide range of particles ranging from silt to clay could be selected for mound building. Clay enrichment in termite mounds is related to translocated mica and smectite or saprolitic materials from the subsoil [50]. According to Jouquet et al. [5], high clay content in termite mounds could arise from the dissolution of plant matter, while resilication is also another process, particularly for smectite.Except at the bottom slope, the studied mounds were predominantly sandy clay loam in texture, and termite mound soils were not different from matched reference pedons, corroborating the findings of Freitas et al. [3]. According to Abe and Wakatsuki [49], the clay mineralogy of the termite mound soil is also very similar to adjacent bulk soils. A contrasting mound soil texture, as with the pedon D02, implies that termites mine and translocate soil particles depending on the nature of the subsoil. In addition, the silt-to-clay ratio of termite mounds ranged from 0.37 to 0.61 which was comparable to the ratio obtained from reference pedons (Table 4). This supports previous studies [50, 51] and indicates that termites relocate fine particles from the subsurface of the respective reference soils across toposequences.


## 3.4. Chemical Properties of Termite-Mediated Soils

Soil pH (1 : 2.5 H2O) varied across the toposequences (Table 4), which could be attributed to the leaching of cations from the upslope and the subsequent accumulation at the nearly level positions [52]. Moderately acidic pH increased with depth in pedon D01 and became strongly acidic at the subsurface of pedon D02, while it was uniformly distributed in pedon D03. The soil pH was neutral at the surface, slightly increased with depth in pedon M01, and changed to moderately alkaline in pedon M02. Apart from pedon M02, the acidic soil surface could be related to the leaching of basic cations, while limited leaching due to herbaceous vegetation probably maintained a near neutral soil reaction in pedon M01, whereas the alkalinity of M02 could be attributed to the nature of the parent material. In general, the comparable soil pH within the pedons likely indicates slow weathering and soil development. The soil pH values of termite mounds ranged from slightly acidic to neutral which were not similar to the respective pedons D01 and D02 but similar to the subsurface horizons of the pedons M01 and M02 (Table 4). The soil pH of mounds depends on the corresponding pH values of the reference soil at a given topographical position. This is consistent with the findings of Sarcinelli et al. [32]. According to Bera et al. [7], the pH of termite mounds is attributed to cation-clay accumulation and the rate of OM decomposition in the mounds.While no carbonate was present in all other soils, the soils in the pedonM02 invariably exhibited strong effervescence with dilute HCl (10%) and showed calcium carbonate that rapidly increased with depth (Table 4). Termite mounds at the same position had CCE that was 2.5 times higher than the respective surface soil and comparable to the subsurface of the reference pedon. The accumulation of carbonate within a pedon could be inherited from parent materials and other associated rocks. The CCE-depth relationship also supports the premise that it is geological rather than pedogenic since weathering of the bedrock results in substantial Ca2+ [41]. According to Buss et al. [53], the carbonate incidence near the surface soil suggests a limited percolation of water to leach carbonate. The distinct features of CCE in mounds indicate that termite pedoturbation concentrates carbonate and relays it to bulk soils. This result is consistent with the observation of Mcauliffe et al. [54] who reported the enrichment of mounds with carbonate.Soil EC content showed an irregular pattern across topographical positions owing to leaching and accumulation of basic cations (Table4). Apart from pedon M02, the EC in the horizons of the pedons was below 2.34 mS cm−1. For the pedon M02, the EC value of 8.66 mS cm−1 in the surface soil decreased gradually with depth. In general, negligible EC differences in the soils of most pedons indicate limited eluviation and illuviation processes, while the amounts in the pedon M02 could be related to the substantial basic cations in the soils of the pedon. Compared to the reference pedon, the average EC of mounds was almost seven times higher than the maximum value for soils in the pedons D01 and M01 and 43 times higher in the pedon D02 while comparable in the pedon M02. By and large, the higher EC of termite mounds compared to bulk soils could be attributed to more soluble ions in the biogenic structure [55].The soil OC varied consistently along the toposequences (Table4). This corroborates with the findings of Sarcinelli et al. [32] which suggest that similar compositions of termite foraging material result in comparable OC content across topographical positions. Except for the pedon D03 which had a wetting-front pattern, the OC gradually decreased with depth. The OC of mounds at the summit and back slope exceeded the higher content of the surface horizons by more than 70%. However, the OC content of the mounds was lower than that of the surface bulk soils of the pedons M01 and M02. While wetting-front OC trends might be attributed to slow translocation, the gradual decreasing pattern probably indicates the accumulation of OC through grass decay. Based on the OC-to-clay index stated by Merante et al. [56], the high to moderate OC content in pedon M02 could also be associated with a complex OC with clay content, which could lead to C stabilization. Regarding the OC levels of the mounds, the result is consistent with the observation of Obi et al. [57] and demonstrates topographic effects. In addition to the high mineralization rate in a semiarid area, the fact that fungus-growing termites mineralize about 20% of the total OC [11] probably results in a low OC content.The available P was less than 10.69 mg·kg−1 across the studied pedons and irregularly distributed across toposequences (Table 4). The distribution pattern of P showed accumulation with depth in the pedons D01, D03, and M02, while its value gradually decreased in the pedons D02 and M01. Compared to the reference pedon, the average P content of the mounds was higher by 51%, 163%, and 78% than the maximum values of the soil horizons in the pedons D02, M01, and M02, respectively. In general, low available P could be attributed to insoluble Ca phosphates comprising the calcareous materials in the pedon M02, while the low levels in the other pedons might be due to sorption by sesquioxides. According to Dinssa and Elias [42], a decreasing P-trend with depth could be associated with reduced OM and P fixation. The highly available P of termite mounds could be related to the incorporation of OM into mounds [4]. However, Bera et al. [7] reported fewer contents in mounds arguing that P content depends on the termite feeding habit and the soil materials. The P content of the mound also depends on the amount of clay, pH, types, and termite species involved in mound formations [11]. In our case, the viable explanation for the highly available P content of mounds is probably related to the soil pH nearing neutral reaction, clay content, and long-term accumulation of OM leading to humification which in turn favors the release of P.The CEC of the soils varied irregularly across the toposequences (Table4). In pedons D01 and D02, CEC had an irregular pattern with depth. With a rapidly changing pattern, the pedons D03 and M01 had increasing CEC values with depth, while pedon D03 had substantially high CEC in the lowest horizon. The CEC gradually decreased with depth and accumulated in the lowest horizon of the pedon M02. A low CEC in the surface soil could be associated with a low OM content, while a moderate to high CEC in the subsurface could result from clay accumulation. A distinctly high CEC in the pedon D03 could be due to clay synthesis (r > 0.99∗∗, not shown here) and the accumulation of Ca2+ and Mg2+ (r = 0.99∗∗). On the other hand, the considerable CEC in the pedon M02 could be associated with substantial clay and Ca2+ in the soils [41]. The CEC values of the mounds on the summit and back slope were three times, while that of the mounds at the foot slope and bottom slope were two-fold higher than the maximum CEC in the respective horizons of the reference pedons. The higher CEC of termite mounds might be independent of OM (r = 0.19 ns), and it is rather related to the presence of 2 : 1 clay. This is because the CEC of both soil and clay is higher than what would be expected for soils dominated by kaolinite, which suggests the presence of montmorillonite clay in the soil of termite mounds and related to the subsoil of respective pedons. This corroborates with the findings of Abe and Wakatsuki [49] who observed minor modification of clay mineralogy in Macrotermes mounds compared to their respective soils. Consequently, this clay probably explains the high CEC of termite mounds.While Ca2+, Mg2+, and K+ showed an increasing trend, Na+ exhibited decreasing patterns across topographical positions (Table 5). The increasing pattern of Ca2+ and Mg2+ down the slope could be due to lateral movement. Exchangeable Ca2+ was less than 10 cmolc kg−1 in all pedons except for pedon M02 and a distinctly high value in the lowermost horizon of pedon D03. Both Ca2+ and Mg2+ had an irregular pattern with depth in the pedons D01 and D02 and M02, rapidly changed with depth in the pedon D03, and showed accumulation in the subsurface of the pedon M01. The high Ca2+ in the pedon D03 could be related to the CaCO3 accumulation [53] and that of the pedon M02 could be attributed to the weathering of carbonate-enriched parent materials [41]. Exchangeable K+ gradually increased in the pedons D01 and D02, while the other pedons gradually decreased with depth. In all pedons, Na+ had an irregular decreasing trend with depth except within pedon D03 which had some enrichment. The increase of K+ with depth in the pedons D01 and D02 could be related to the leaching of cations [40]. Relatively, the high K+ in the pedon D03 is probably due to the accumulation of mica at that position, while the Na+ accumulation in the lowest horizon of this pedon may be due to its greater mobility. The Ca > Mg > K > Na order in the studied pedons might be due to the release of these ions by weathering before the rocks form. The major cations that dominated the exchange complex of the pedons are consistent with the findings by Beyene [58].Table 5
Exchangeable cations and available micronutrients of the termite mounds (TM,n = 3, ± standard deviation) and reference profiles across toposequences in Ilala-Sara and Silala, southeast Ethiopia.
HorizonsSampling depth (cm)Exchangeable cationsAvailable micronutrientsCa2+Mg2+K+Na+FeMnZnCucmolc kg−1mg kg−1Profile D01, Ilala-Sara, ∼ gently undulating summitA0–120.470.400.190.147.0810.300.940.50AB12–300.910.770.190.1210.539.271.390.71Bw30–730.650.480.160.135.775.821.390.64BC73–1401.370.460.260.116.515.391.220.57Cr140–200+1.010.750.270.1210.951.750.140.36TM6.61 ± 0.640.47 ± 0.120.13 ± 0.030.13 ± 0.04————Profile D02, Ilala-Sara site, ∼ gently undulating back slopeA0–71.680.510.110.1511.275.360.270.38AB7–951.210.610.150.159.466.730.090.31Bw95–1401.420.460.120.162.891.020.790.33BC140–1701.240.690.160.1311.331.820.420.36Cr170–200+1.310.530.190.149.442.020.220.36TM19.15 ± 1.310.72 ± 0.140.13 ± 0.020.10 ± 0.00————Profile D03, Ilala-Sara site, ∼ nearly levelled toe slopeA0–101.781.080.380.1013.659.950.520.40AB10–401.671.130.210.0618.2015.980.530.45Bg40–701.190.870.290.0725.385.470.230.54Bt70–200+12.223.200.220.1920.081.940.630.97Profile M01, Silala site, ∼ very gentle foot slopeA0–103.930.330.180.1532.861.932.070.72AB10–552.360.250.150.1242.609.261.741.23Bt55–904.351.300.170.1737.286.691.581.13Bo90–200+5.531.720.150.1431.283.431.240.78TM8.69 ± 0.791.17 ± 0.270.23 ± 0.020.14 ± 0.04————Profile M02, Silala site, ∼ levelled bottom slopeA0–1851.141.500.150.1330.705.191.870.50AB18–4251.470.700.170.1720.262.751.320.42Bk42–7054.651.710.150.1229.325.111.720.43Ck70–12047.124.480.150.1021.304.490.290.46Ckm120–200+31.464.750.100.0927.246.111.380.41TM70.24 ± 2.153.38 ± 0.120.23 ± 0.030.07 ± 0.03————The mounds at the summit and back slope had 4 to 10 times higher Ca2+, while the mounds at the bottom slope had 57 times higher Ca2+ than the maximum value in the respective reference pedon (Table 5). On the other hand, the Mg2+ contents of the mounds were lower compared to that of the surface soils but comparable to the levels in the subsurface horizon of the reference pedons. The K+ content of mounds decreased up to 52% compared to the maximum values in the soils of the reference pedons D01 and D02, while the K+ content of the mounds was higher by more than 28% in the pedons M01 and M02. The exchangeable Na+ of the mounds was decreased by 7% to 59% compared to the maximum values in the reference pedons. While termite pedoturbation did not affect the Na+ and Mg2+ across the toposequences, their effects were evident for Ca2+ at all locations and K+ at the foot slope and bottom slope positions. In support of this study, Obi et al. [57] also considered Ca2+, Mg2+, and K+ as indicators of pedoturbation.


## 3.5. Available Micronutrients

While Zn and Cu decreased down the slope, Fe and Mn showed irregular patterns across toposequences (Table5). Consistent with Hailu et al. [43], the patterns of Fe and Mn were similar at any given topographical position because of their similar chemical behavior. The micronutrient contents were in the order of Fe > Mn > Zn > Cu. According to Zhu et al. [59], the dominance of Fe at any given topographical position might be due to the dissolution of metal oxide in the clay particles which in turn strongly depends on the slope position while the availability of Zn and Cu related to desorption from clay, carbonate, and OM surface. Available Fe, Mn, and Zn contents showed an irregular pattern with depth in the pedons D01, D02, and M02, probably owing to termite pedoturbation while accumulating with depth in pedon D03. While Fe and Mn accumulated just below the surface layer, Zn gradually decreased with depth in the pedon M01. Available Cu showed a gradual decrease with depth in the pedons D01, D02, and M02, while it gradually increased in the pedons D03 and M01.


## 3.6. Pedogenesis and Classification of the Soils

The subsurface horizon of the pedonsD01 and D02 showed an incipient pedogenetic alteration, with half of the original rock structure being obliterated. Additionally, the horizon thickness and texture of the soils qualify them for a cambic horizon. The cambic horizons were also specifically identified by redder hues (2.5 years) and higher chroma (>6) on the surface than the underlying horizons. On the other hand, sufficiently intense reduction and segregation of iron which resulted in low chroma were indicative of the identification of the cambic horizon in the pedon D03. Prominent clay accumulation in the subsoil horizons of the pedon D03 qualified argic diagnostic horizon. The pedon M01 was differentiated by a redder hue, higher chroma, and an accumulation of clay and sesquioxides. The moist color changes from reddish-brown in the upper to dark reddish-brown at the subsurface horizon substantiating clay illuviation for identification of the argic horizon. Termite pedoturbation [13], shrubs, perennials, and grasses probably resulted in clay accumulation in this pedon. A polyhedral structure in this pedon did not meet the diagnostic features of a nitic horizon. The subsurface horizon of the pedon M02 had more than 15% CCE in the fine Earth fraction which meets the diagnostic criteria for a calcic horizon. According to Macedo et al. [60], the absence of secondary carbonate accumulation may be due to termite pedoturbation. Based on the above diagnostic horizons and using established criteria from IUSS Working Group WRB [18], the pedons D01 and D02 were classified as Cambisols. Argic horizons accompanied by CECs of clay greater than 24 cmolc kg−1 in the pedons D03 and M01 differentiated the soils into Luvisols, whereas the pedon M02 was identified as Calcisols. The principal and supplementary qualifiers are summarized in Table 6.Table 6
Descriptions of diagnostic horizons and soil classification based on the WRB system.
Ilala-Sara, Dugda DawaSilala, MiyoPedon,D01Pedon,D02Pedon,D03Pedon,M01Pedon,M02Diagnostic horizonCambic horizonCambic horizonArgic horizonArgic horizonCalcic horizonIUSS working group WRB [18]Hyperdystric endoskeletic cambisols (clayic and humic)Dystric skeletic cambisols (loamic and humic)Gleyic luvisols (clayic, neocambic, differentic, epidystric, and humic)Chromic luvisols (clayic, epidystric, and humic)Skeletic cambic calcisols (loamic and humic)


## 3.7. Pedogenic Processes

The soils in the surface layers of all pedons showed the feature of termite pedoturbation as traced by granular or subangular blocky structures and gradual or diffuse horizon boundaries except for soils in pedonD03. In addition, incipient pedogenic processes, as shown by little alteration and very few fine root distributions, might be some of the soil-forming processes in the pedons D01 and D02. As stated by Borden et al. [52], the loss of cations through downslope translocation also probably promotes desilication in these two pedons. The evidence of gleization was observed in the pedon D03, but cyclic drying and wetting in the surface and the remains of moisture in the subsoil favored hydrolysis which subsequently derived clay formation. In this pedon, the absence of mottling suggests minimal current pedogenetic activities other than gleying and illuviation processes. According to Bonifacio et al. [44], gleyic properties indicate strong weathering of the iron-bearing minerals probably as evidenced by the depletion of available Fe on the surface of the pedon. In the pedon M01, desilication, braunification, rubification, ferrugation, leucinization, and illuviation might be the most important pedogenic processes. Calcification was the dominant process in the pedon M02 while humification dispersed to a certain extent throughout the solum. Following Deressa et al. [36], there was a textural differentiation in examined pedons except for the pedon M02 which showed a clay contrast index of 0.87 indicating a lower textural gradient. In addition to termite pedoturbation, calcification processes result in homogenous clay distribution within the pedon M02.


## 4. Conclusions

Termite pedoturbation promotes soil genesis by modifying soil morphology, sorting particles, and altering physicochemical properties. However, the magnitude of their impact is, by and large, specific to topographical positions. The studied soil properties also show topographic effects exhibited by various degrees of variation along the toposequences where gently sloped positions had the highest clay, K+, and Na+, while sand, pH, OC, Ca2+, and Mg2+ were high in nearly level positions. This study also showed the reference subsoils are mined for fine particles, and thus, there is a slight modification to B horizons across the toposequences. The soils of the study sites are grouped into Cambisols, Luvisols, and Calcisols. Pedogenic processes that formed Cambisols on the summit and back slope are less active processes complemented by termite pedoturbation. Luvisols are formed by gleying and clay synthesis at the toe slope where termites are absent or by illuviation processes and termite pedoturbation on the foot slope. Calcisols occur to a lesser extent on nearly level land, while calcification and termite pedoturbation mainly operated during pedogenesis. Based on the results of this study, termite pedoturbation and topography should be carefully considered during the classification and management of semiarid soils. Although this study marked the influence of termite activity and topography, the precise effects are warranted following micropedological evidence.



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*Source: 7150907-2023-01-27.xml*