Effects of termites on the physical and chemical properties of the acid sandy soils of southern Nigeria

Abstract

This study was aimed at characterizing the effects of the activity of termites of the genus Nasutitermes on the physico‐chemical properties of the acid sandy soils of southern Nigeria. Selected morphological properties of the termite mounds were measured in the field. Outside portions of the termite mound and surface (0–15 cm) soil were collected and analyzed for some physical and chemical properties. Results obtained showed a density of 112 mounds ha^‐1 with average height of 0.85 m. There were significantly higher proportions of clay, silt, and organic carbon, and higher pH, exchangeable potassium (K), calcium (Ca), magnesium (Mg), available phosphorus (P), effective cation exchange capacity and base saturation in the mounds of the Nasutitermes than in the surrounding topsoil. Mounds of Nasutitermes termites, if returned to the soil, could improve the properties of the soil in areas where termites occur in large numbers.     

Modification of soils in Nigerian savanna by soil-feeding Cubitermes (isoptera,termitidae)

Soil-feeding termites ingest humified, organic-rich soil. The soil faeces are used for nest construction and mounds of two species of Cubitermes contained more soil, clay, exchangeable Ca and Mg, available P, total N and organic C than adjacent topsoil. Available P increased by 1.4–6.0 times. Mounds of a plant-debris feeding termite, Trinervitermes, contained significantly more of these fractions, with the exception of available P, than adjacent topsoil. The modification of Trinervitermes mounds by Cubitermes resulted in a 2-fold increase in available P, whereas organic C remained the same and N increased by 1.5-times. The relatively large increase in available P resulting from soil feeding termites could be attributed to the high pH regime in their hind-guts.     

High performance liquid chromatography studies on the polysaccharides in the walls of the mounds of two species of termite in Senegal,Cubitermes oculatus and Macrotermes subhyalinus: their origin and contribution to structural stability

 The origin, nature and quantity of polysaccharides in the walls of the epigeal mounds of a species of soil-feeding termite, Cubitermes oculatus, and a fungus-growing termite, Macrotermes subhyalinus, found in Senegal, and of soil not considered to be under the influence of termites, were studied to obtain a clearer picture of the structural stability of these materials. The compounds were extractedand analysed by high performance liquid chromatography. We found that the walls of mounds made by soil-feeding species were very rich in sugars soluble in aqueous acid or hot water. Most of the sugars originated from cellulose and hemicellulose, and only a small proportion from microorganisms. There were also significant amounts of stachyose in the mound walls and in the reference soil. This sugar was probably formed by the surrounding vegetation, which was mainly leguminous crops. Comparison of the mineral and organic-mineral particle sizes of samples confirmed that the walls of soil-feeding termite mounds where there is the greatest redistribution of clay have the best aggregating capacity. The results therefore show that the polysaccharides in mound walls of soil-feeding termites are mostly of plant origin. Their influence on the stability of these structure is discussed. The walls of fungus-growing termite mounds contain little organic matter and hence low levels of polysaccharides, which are mainly of plant origin. Received: 19 July 1999     

Some effects of mound-building termites on the soils of a semi-arid area of Kenya

The nutrient status of soils from Macrotermes termite mounds, in and around Kajiado District, reflects that of the subsoils more closely than in some other studies, notably Watson (1977). Consequently the growth of vegetation on and around mounds was not noticeably enhanced, except in grasslands and in higher rainfall areas. There was no evidence that mounds acted as wicks. Mound soils contained greater concentrations of calcium, magnesium and potassium than subsoils in almost all cases where subsoil values were low, suggesting that the termites’ activities enhanced the nutrient status of mound soils. The nutrients were probably derived from their food, and the effects increased in some cases by their selection of clay on clay-poor soils. Nevertheless, the overall effects of Macrotermes on rangeland soils are probably small. The distribution of Macrotermes is remarkably unaffected by soil types; they construct mounds on almost all soils except those consisting largely of montmorillonite.     

Termite mounds in relation to the surrounding soils in the forest and derived savanna zones of southeastern Nigeria

Some physical and chemical properties of the two common termite mounds in southeastern Nigeria, Macrotermes (MM) and Cubitermes (CM) mounds, were compared and their relationships with the surrounding top and subsoils investigated. Percentage sand, silt, pH, calcium, magnesium, potassium, sodium, iron, and organic carbon were higher and clay and penetrometer resistance lower in the CM than the MM. Sand, silt, organic carbon, and calcium decreased and clay increased from the forest to the derived savanna in the CM, but no clear trend was shown with the other properties in both the CM and the MM. Most of the nutrients were higher in the CM than in the adjacent top or subsoil but the reverse was the case with the MM. More nutrients are associated with the inorganic fractions of the soil than with the organic carbon in the mounds and the soils surrounding them. The Ca:Mg ratio in the mounds and the top and subsoils adjacent to them was low for most crops but the K:Mg ratio was mostly adequate. Because of these differences in properties and sizes of the mounds, different management strategies are recommended for them and the soils around them.     

Identifying potential management zones in a layered soil using several sources of ancillary information

Variation in soil texture has a profound effect on soil management, especially in texturally complex soils such as the polder soils of Belgium. The conventional point sampling approach requires high sampling intensity to take into account such spatial variation. In this study we investigated the use of two ancillary variables for the detailed mapping of soil texture and subsequent delineation of potential management zones for site‐specific management. In an 11.5 ha arable field in the polder area, the apparent electrical conductivity (EC_a) was measured with an EM38DD electromagnetic induction instrument. The geometric mean values of the EC_a measured in both vertical and horizontal orientations strongly correlated with the more heterogeneous subsoil clay content (r = 0.83), but the correlation was weaker with the homogenous topsoil clay content (r = 0.40). The gravimetric water content at wilting point (θg_{(?1.5 MPa)}) correlated very well (r = 0.96) with the topsoil clay content. Thus maps of topsoil and subsoil clay contents were obtained from 63 clay analyses supplemented with 117θg_{(?1.5 MPa)} and 4048EC_a measurements, respectively, using standardized ordinary cokriging. Three potential management zones were identified based on the spatial variation of both top and subsoil clay contents. The influence of subsoil textural variation on crop behaviour was illustrated by an aerial image, confirming the reliability of the results from the small number of primary samples.     

Lime placement on subsoil as a strategy to reduce phosphorus leaching from agricultural soils

Leaching of phosphorus (P) from agricultural land is a major contributor to eutrophication of surface waters in many countries, and effective mitigation options to reduce P in leachate are needed. In this study, intact columns (0.77 m deep) of subsoil from three Swedish agricultural soils (one sand and two clay) were used to examine whether placing quicklime (calcium oxide, CaO) on the subsoil could reduce P leaching over a 3‐yr period. Leaching of particulate P (PP) was significantly less from clay soil columns with lime than from clay soil columns without (P < 0.001 and P < 0.05, respectively), with a relative reduction of 49 and 51% in the two soils. Leaching of dissolved reactive P (DRP) was less from sand columns with lime than from sand columns without, although not significantly so due to large variation in P leaching between columns. These results indicate that placement of lime on subsoil has potential to reduce P leaching, especially of PP from clay soils. However, more studies including both topsoil and subsoil and a range of soil types are needed to assess the full potential of this P mitigation option.     

Nutrient storage in soils and nests of mound-building <Emphasis Type="Italic">Trinervitermes</Emphasis> termites in Central Burkina Faso: consequences for soil fertility

Nest structures of two termite species (Trinervitermes spp.) with epigeal (above-ground) mounds were analyzed to compare their nutrient status with that of adjacent soils. To take into account soil variability, the observations and samplings were made in three toposequences of different and representative West African savanna soils. The data showed the high degree of adaptation of these termite species to a large range of soil types and environments. Mounds of Trinervitermes geminatus and Trinervitermes trinervius, both grass-feeders, contained more clay, organic matter (OM), and exchangeable cations than the surrounding surface layer soil. The storage of OM and exchangeable cations was determined for T. geminatus nests and compared to the surrounding soil. Despite substantial nutrient storage in mounds, its total weight appeared low when compared to the nutrient storage in the surrounding 0–15 cm of soil surface layer. This illustrates how contradictory points of view on the use of termite mounds in agriculture need to be clarified using a classical approach that takes into account data by species; and this also evaluates the contribution of different termite mounds to nutrient fluxes and storage and the exact stocking rate of living mounds.     

Impact of subterranean fungus-growing termites (Isoptera, Macrotermitiane) on chosen soil properties in a West African savanna

Fungus-growing termites (Isoptera, Macrotermitinae) play an important role in tropical ecosystems in modifying soil physical properties. Most of the literature regarding the impact of termites on soil properties refers to termite epigeous mounds. In spite of their abundance and activity in African savannas, few studies deal with the properties of underground nest structures (fungus-comb chambers) built by subterranean Macrotermitinae termites. We tested whether these termites significantly modify the soil physico-chemical properties within their nests in a humid tropical savanna and whether these effects are different for two termite species with differing building behaviour. Termite-worked soil material was collected from fungus-comb chamber walls of two widespread species: Ancistrotermes cavithorax, which builds diffuse and ephemeral nests and Odontotermes nr pauperans, which most often builds concentrated and permanent nests for a comparatively much longer period of time. Neither species influenced soil pH but both significantly modified soil texture and C-N content in their nest structures. A strong impact on clay-particle size was also detected but no significant differences in clay mineralogy. Thus Odontotermes has a greater effect on soil properties, that could be explained by its building behaviour and the concentration in space of its nest units. Therefore, spatial pattern and life-span of fungus-comb chambers should be an important parameter to be considered in the functional role of subterranean Macrotermitinae termites in the savanna.     

Accumulation of free oxyhydroxides in termite (Macrotermes bellicosus (Smeathman)) mounds and the implications for their dynamics in an African savannah Ultisol

The present study aimed to assess the dynamics of oxyhydroxides via termite mounds in a tropical savannah of Central Nigeria, where the soils often contain oxyhydroxides as a major component of soil minerals. To this end, the quantities of oxyhydroxides stored in mounds built by Macrotermes bellicosus (Smeathman) were compared to those stored in surface (Ap1) soils, and their turnover rates were estimated. Both the mound wall and nest of M. bellicosus were enriched two- to 10-fold with acidified ammonium oxalate soluble iron (Fe_o) and aluminum (Al_o) and dithionite-citrate-bicarbonate (DCB) soluble iron (Fe_d) and aluminum (Al_d) relative to the adjacent surface soil horizon. These oxyhydroxide contents were positively correlated with the clay content (P < 0.05), suggesting that M. bellicosus preferentially used silicate clay-associated oxyhydroxides for mound construction. The Fe_d, Al_d and DCB-soluble manganese (Mn_d) preserved in the M. bellicosus mounds ran up to 112 ± 25.6, 5.72 ± 1.41 and 2.17 ± 0.68 kg ha^?1, accounting for 1.91 ± 0.23%, 1.00 ± 0.60% and 0.35 ± 0.09% of the total amount stored in the surface soil horizon, respectively. Furthermore, the estimated turnover rates of Fe_d, Al_d and Mn_d were 6.6, 0.33 and 0.14 kg ha^?1 year^?1, respectively. These findings suggest that the mound-building termites significantly impacted the dynamics of free oxyhydroxides in an African savannah soil.     

Interacting effects of topsoil water and nitrogen supply on subsoil aluminium toxicity

Abstract

The interacting effects between topsoil water supply, nitrogen (N) placement and subsoil aluminum (Al) toxicity on wheat growth were studied in two split‐root pot experiments. The native nitrate‐N (NO₃‐N) in the topsoil used in each experiment differed and were designated as high (3706 μM) and low (687 μM) for experiments one and two, respectively. Wheat was grown in pots that enabled the root system to be split so that half of the roots were in topsoil and the other half were in subsoils containing varying concentrations of soluble Al. Treatments were imposed which varied the supply of water to the topsoil (either ‘wet’ or ‘dry'). Placement of applied N in either the topsoil or subsoil had little effect on either shoot or root fresh weight, or on the length of roots produced in the subsoil section of the split pots. When water supply to the topsoil was decreased, both shoot and root growth of wheat declined and the yield decrease increased with subsoil Al. In the high‐N experiment, wheat grown in the low Al subsoil with the high native soluble subsoil (NO₃ (3002 μM) was able to exploit the N and subsoil water, hence both shoot and root growth increased considerably in comparison to shoot and root growth of wheat grown in soils containing higher concentrations of subsoil Al. When the native NO₃ was lower (i.e. the low‐N experiment) inadequate root proliferation restricted the ability of plants to use subsoil N and water irrespective of subsoil Al. The results from this study suggest that wheat, grown on yellow earths with Al‐toxic subsoils, will suffer yield reductions when the topsoil dries out (e.g. in the spring when winter rainfall ceases) because subsoil reserves of water and nitrogen are under utilised.     

Physicochemical and morphological properties of termite (Macrotermes bellicosus) mounds and surrounding pedons on a toposequence of an inland valley in the southern Guinea savanna zone of Nigeria

Abstract

Termites play a significant role in soil-forming processes of the tropics. The influence of termites on pedogenesis as affected by the toposequence, however, has rarely been explored. We investigated the soil physicochemical and morphological characteristics of epigeal mounds constructed by Macrotermes bellicosus (Smethman) compared with those of surrounding pedons along a toposequence (bottom, fringe and upland sites) of an inland valley in central Nigeria. The physicochemical and morphological properties of the mound soils varied according to structural units but were generally different from those of the adjacent pedons. The differences included finer texture, higher electrical conductivity, total N, exchangeable bases (Ca, Mg and K) and effective cation exchange capacity and lower C/N ratio and exchange acidity in the mound than the pedon at each toposequence position. This tendency to modify the soil properties was more prominent in the nest body where the termites actually live, that is, in the hives, royal cell and base-plate, than in the soils below the nest and the other mound parts, that is, the external wall, internal wall and pillars. We found this trend to a greater or lesser degree at all toposequence positions. Our findings suggest that: (1) M.?bellicosus can manipulate the mound soils according to functional applications of structure units or environmental requirements for its livelihood, regardless of local soils; (2) M.?bellicosus makes ecological patches (hot spots) at all toposequence positions in the same measure; (3) the influence of M.?bellicosus on the pedogenesis is reduced in the lowlands compared with the uplands because the number and volume of the mounds were substantially lower in the bottom and fringe sites compared with the upland site.     

Effect of different termite feeding groups on P sorption and P availability in African and South American savannas

Nest structures of six termite species, four with epigeous (above-ground) and two with subterranean nests were analysed to find out how their building and feeding habits could be related to their nests phosphorus status compared with control soils. Termite nest structure was found to affect significantly the P status in savanna soils: mounds of the African Trinervitermes geminatus and the South American Nasutitermes ephratae (both grass-feeders) displayed a greater amount of available P, especially in the inner part of the nest, than the surrounding soil. The abundant quantities of dead grass material stored in the mound can explain the available soil P increase. A similar increase in P availability was also found for the soil-feeder Cubitermes severus. In mounds of Macrotermes bellicosus, on the other hand, there was a drastic increase in P sorption (and a corresponding decrease in available P) compared to adjacent soils, which was attributed to the building strategy of this species. M. bellicosus selected clay from subsoil to build its nest structure. The data obtained for the subterranean species Ancistrotermes cavithorax and Microtermes toumodiensis indicated also that there is an increase in P sorption in mounds when compared with associated topsoils. Consequently, the nest structures of only certain termite species should be considered, and utilised, as a soil amendment in place of fertilisers. This impact on the P cycle in savannas seems to be related to the termite feeding status and to the type of material utilised in nest building. This should be taken into account before using termite nest material in soil fertility status improvement.     

Enhancing pedotransfer functions with environmental data for estimating bulk density and effective cation exchange capacity in a data‐sparse situation

Soil bulk density (BD) and effective cation exchange capacity (ECEC) are among the most important soil properties required for crop growth and environmental management. This study aimed to explore the combination of soil and environmental data in developing pedotransfer functions (PTFs) for BD and ECEC. Multiple linear regression (MLR) and random forest model (RFM) were employed in developing PTFs using three different data sets: soil data (PTF‐1), environmental data (PTF‐2) and the combination of soil and environmental data (PTF‐3). In developing the PTFs, three depth increments were also considered: all depth, topsoil (<0.40 m) and subsoil (>0.40 m). Results showed that PTF‐3 (R²; 0.29–0.69) outperformed both PTF‐1 (R²; 0.11–0.18) and PTF‐2 (R²; 0.22–0.59) in BD estimation. However, for ECEC estimation, PTF‐3 (R²; 0.61–0.86) performed comparably as PTF‐1 (R²; 0.58–0.76) with both PTFs out‐performing PTF‐2 (R²; 0.30–0.71). Also, grouping of data into different soil depth increments improves the estimation of BD with PTFs (especially PTF‐2 and PTF‐3) performing better at subsoils than topsoils. Generally, the most important predictors of BD are sand, silt, elevation, rainfall, temperature for estimation at topsoil while EVI, elevation, temperature and clay are the most important BD predictors in the subsoil. Also, clay, sand, pH, rainfall and SOC are the most important predictors of ECEC in the topsoil while pH, sand, clay, temperature and rainfall are the most important predictors of ECEC in the subsoil. Findings are important for overcoming the challenges of building national soil databases for large‐scale modelling in most data‐sparse countries, especially in the sub‐Saharan Africa (SSA).     

The physical and chemical composition of mounds of Macrotermes carbonarius (Hagen) (Termitidae, Macrotermitinae), in Penang, Malaysia

The mound of the fungus-growing termite Macrotermes carbonarius (Hagen) contained significantly larger proportions of clay (P<0.05) than the adjacent soil. The largest clay content was found in the brood chamber. Concentrations of organic carbon, nitrogen, potassium and calcium in various parts of the mound were enriched compared to the adjacent soil at 10–40 cm depth from which they might have been derived. The activities of M. carbonarius did not significantly alter the pH or the bulk density of the mound soil compared to the adjacent topsoil.     

Effect of termites on clay minerals in tropical soils: fungus-growing termites as weathering agents

Termites of the subfamily Macrotermitinae play an important role in tropical ecosystems: they modify the soil's physical properties and thereby make food available for other organisms. Clay is important in the architecture of Macrotermitinae termite nests, and it has been postulated that termites could modify the mineralogical properties of some clays. We have tested this hypothesis of clay transformation by termites in the laboratory under controlled conditions, using Odontotermes nr. pauperans termite species, one of the main fungus‐growing species at Lamto Research Station (Côte d'Ivoire). Soil handled by termites in nest building was saturated with SrCl₂, glycol or KCl and afterwards heated at 250°C for X‐ray diffraction analyses. Termite handling led to an increase in the expandable layers of the component clay minerals. Heating and saturation by potassium of modified clays did not close the newly formed expandable clay layers. However, differences occurred between parts of the constructions built by termites, and the clays can be ranked according to their degree of alteration in the following order: unhandled soils < galleries < chamber walls. Consequently, termites can be seen as weathering agents of clay minerals, as previously shown for micro‐organisms and plants.     

Nutrient storage in termite (Macrotermes bellicosus) mounds and the implications for nutrient dynamics in a tropical savanna Ultisol

The role of mounds of the fungus-growing termite Macrotermes bellicosus (Smeathman) in nutrient recycling in a highly weathered and nutrient-depleted tropical red earth (Ultisol) of the Nigerian savanna was examined by measuring stored amounts of selected nutrients and estimating their rates of turnover via the mounds. A study plot (4?ha) with a representative termite population density (1.5?mounds?ha^?1) and size (3.7?±?0.4?m in height, 2.4?±?0.2?m in basal diameter) of M. bellicosus mounds was selected. The mounds were found to contain soil mass of 9249?±?2371?kg?ha^?1, composed of 7502?±?1934?kg?ha^?1 of mound wall and 1747?±?440?kg?ha^?1 of nest body. Significant nutrient enrichment, compared to the neighboring topmost soil (Ap1 horizon: 0–16?cm), was observed in the nest body for total nitrogen (N) and exchangeable calcium (Ca), magnesium (Mg) and potassium (K), and in the mound wall for exchangeable K only. In contrast, available (Bray-1) phosphorus (P) content was found to be lower in both the mound wall and the nest body than in the adjacent topmost soil horizon. Consequently, the mounds formed by M. bellicosus contained 1.71?±?0.62?kg?ha^?1 of total N, 0.004?±?0.003?kg?ha^?1 of available P, 3.23?±?0.81?kg?ha^?1 of exchangeable Ca, 1.11?±?0.22?kg?ha^?1 of exchangeable Mg and 0.79?±?0.21?kg?ha^?1 of exchangeable K. However, with the exception of exchangeable K (1.2%), these nutrients amounted to less than 0.5% of those found in the topmost soil horizon. The soil nutrient turnover rate via M. bellicosus mounds was indeed limited, being estimated at 1.72?kg?ha^?1 for organic carbon (C), 0.15?kg?ha^?1 for total N, 0.0004?kg?ha^?1 for available P, 0.15?kg?ha^?1 for exchangeable Ca, 0.05?kg?ha^?1 for exchangeable Mg, and 0.06?kg?ha^?1 for exchangeable K per annum. These findings suggest that the mounds of M. bellicosus, while being enriched with some nutrients to create hot spots of soil nutrients in the vicinity of the mounds, are not a significant reservoir of soil nutrients and are therefore of minor importance for nutrient cycling at the ecosystem scale in the tropical savanna.     

Subsoil rhizosphere modification by chickpea under a dry topsoil: implications for phosphorus acquisition

Chickpea (Cicer arietinum L.) roots exude carboxylates. While chickpea commonly grows where the topsoil dries out during crop growth, the importance of carboxylate exudation by the roots and mobilization of soil P from below the dry topsoil has not been examined. The study investigates the response of carboxylate exudation and soil P mobilization by this crop to subsoil P fertilizer rate. In constructed soil columns in the glasshouse, the P levels (high, low, and nil P) were varied in the well‐watered subsoil (10–30 cm), while a low level of P in the dry topsoil (0–10 cm) was maintained. At flowering, rhizosphere carboxylates and rhizosphere soil from topsoil and subsoil roots were collected separately and analyzed. The concentration of total carboxylates per unit rhizosphere mass in the subsoil was nearly double that of the topsoil. Plants depleted sparingly soluble inorganic P (P_i), NaOH‐P_i, and HCl‐P_i, along with the labile P_i (water soluble and NaHCO₃‐Pi). The P depletion by plants was greater from the subsoil than the topsoil. The study concluded that depletion of sparingly soluble P from the chickpea rhizosphere in the subsoil was linked with the greater levels of carboxylates in the rhizosphere. These findings indicate that chickpea, with its deep rooting pattern, can increase its access to subsoil P when the topsoil dries out during crop growth by subsoil rhizosphere modification.     

Initial soil organic carbon concentration influences the short-term retention of crop-residue carbon in the fine fraction of a heavy clay soil

Among factors controlling decomposition and retention of residue C in soil, effect of initial soil organic C (SOC) concentration remains unclear. We evaluated, under controlled conditions, short-term retention of corn residue C and total soil CO₂ production in C-rich topsoil and C-poor subsoil samples of heavy clay. Topsoil (0–20 cm deep, 31.3 g SOC kg^?1 soil) and subsoil (30–70 cm deep, 4.5 g SOC kg^?1 soil) were mixed separately with ¹³C–¹⁵N-labeled corn (Zea mays L.) residue at rates of 0 to 40 g residue C kg^?1 soil and incubated for 51 days. We measured soil CO₂–C production and the retention of residue C in the whole soil and the fine particle-size fraction (<50 μm). Cumulative C mineralization was always greater in topsoil than subsoil. Whole-soil residue C retention was similar in topsoil and subsoil at rates up to 20 g residue C kg^?1. There was more residue C retained in the fine fraction of topsoil than subsoil at low residue input levels (2.5 and 5 g residue C kg^?1), but the trend was reversed with high residue inputs (20 and 40 g residue C kg^?1). Initial SOC concentration affected residue C retention in the fine fraction but not in the whole soil. At low residue input levels, greater microbial activity in topsoil resulted in greater residue fragmentation and more residue C retained in the fine fraction, compared to the subsoil. At high residue input levels, less residue C accumulated in the fine fraction of topsoil than subsoil likely due to greater C saturation in the topsoil. We conclude that SOC-poor soils receiving high C inputs have greater potential to accumulate C in stable forms than SOC-rich soils.     

Possible influence of termites (Macrotermes bellicosus) on forms and composition of free sesquioxides in tropical soils

There has been less concern about soil mineralogical alteration than about soil physical, chemical and biological changes induced by termite nest-building activity. Furthermore, much less attention has been paid to free sesquioxides than to phyllosilicate minerals. In the present study, we conducted field morphological observations and selective dissolution analysis to characterize free sesquioxides in termite (Macrotermes bellicosus) mounds as compared with surrounding pedons in different toposequence positions, i.e., seasonally flooded valley bottom, hydromorphic fringe and well-drained upland sites. Distinctive redoximorphic features, such as surface yellowish layers on mound structures from the fringe site, indicate possible alteration of iron sesquioxide forms in the mounds due to the transportation of soil from reductive (aquic subsoil) to oxidative (epigeal mound) environments by the nest-building activity of M. bellicosus. On the other hand, the iron-soluble content in the dithionite-citrate-bicarbonate (DCB) system (Fe_d) was generally higher in the mound structures than at the adjacent sub-surface (Ap2) horizon at each toposequence position, while there was less difference in the content of acid ammonium oxalate (AAO) extractable iron (Fe_o) as compared to Fe_d. As a consequence, the iron activity index (Fe_d/Fe_o ratio) was found for the most part to be lower in the mound structures than in the neighboring Ap2 horizon. In addition, the content of Fe_d, AAO-soluble Al (Al_o) and DCB-extractable Al (Al_d) was significantly correlated with clay content in these soils. These findings suggest that M. bellicosus preferentially collects clay particles, probably from the clay-rich subsoils, such as the argillic horizon, which has been formed by the co-migration of phyllosilicate minerals and relatively crystalline sesquioxides. The species then likely incorporates them into the mounds, which induces an increase in the Fe_d content relative to that of Fe_o, resulting in a decreased iron activity index in the mound structures.