Hostname: page-component-848d4c4894-jbqgn Total loading time: 0 Render date: 2024-06-24T17:57:42.696Z Has data issue: false hasContentIssue false

Termitaria enhance soil and forest diversity in Deciduous Dipterocarp Forest, Northern Thailand

Published online by Cambridge University Press:  16 April 2024

Manop Kaewfoo
Doi Chiang Dao Watershed Research Station, Sub-Division of Watershed Research, Division of Conservation and Watershed Management, Department of National Parks, Wildlife and Plant Conservation, Bangkok, 10900 Thailand
Sarayudh Bunyavejchewin
National Parks Wildlife and Plant Conservation Department, Bangkok, Thailand
Dokrak Marod
Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand
Decha Wiwatwittaya
Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand
Ian C. Baillie
School of Water, Energy and Environment, Cranfield University, Cranfield, Bedfordshire MK43 0AL, UK,
Stuart J. Davies
Forest Global Earth Observatory, Natural History Museum, Smithsonian Institution, Washington, DC, USA
Stephen H. Hallett*
School of Water, Energy and Environment, Cranfield University, Cranfield, Bedfordshire MK43 0AL, UK,
Corresponding author: Stephen Henry Hallett; Email:
Rights & Permissions [Opens in a new window]


We characterised the soils and vegetation in 15 sets of four quadrats on and around mounds of Macrotermes annandalei (Isoptera, Macrotermitinae) on a plain of deep dystric clay over limestone in Deciduous Dipterocarp Forest in Northern Thailand. Termites have excavated the mounds from the deep calcareous substrate. The mound soils have darker subsoils, larger contents of clays and exchangeable cations, and higher pH values than the surrounding dystric clay loams. The thickets on the mounds are visually different from the surrounding Deciduous Dipterocarp Forest. They have few dipterocarps and are floristically similar to the regionally important Mixed Deciduous Forest. The clear visual differences are confirmed by floristic similarity, cluster, and canonical correspondence analyses for each of the tree, sapling and seedling size classes. The differences between the mound clays and surrounding red clay loams and the associations between soil and forest types are confirmed by ‘t tests’ and the significant correlations of the soil base status with the main floristic axis of the canonical correspondence analyses. Soil variability due to termites and other agents of pedoturbation can significantly contribute to short-range floristic and structural diversity in some dry tropical forests.

Research Article
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (, which permits unrestricted re-use, distribution and reproduction, provided the original article is properly cited.
© The Author(s), 2024. Published by Cambridge University Press


Spatial and temporal variations in the availability of abiotic resources, such as light, water, nutrients, root aeration and site stability, contribute to the high floristic and structural diversity of tropical forests. The soils of many tropical forests are intensively leached and substantially depleted of labile nutrients, especially P and the exchangeable cationic meso-nutrients K, Ca and Mg (Richards, Reference Richards1996). Some of the variability of nutrient availability in tropical forest soils is systematically associated with lithological heterogeneity in soil parent materials (Baillie et al. Reference Baillie, Ashton, Court, Anderson, Fitzpatrick and Tinsley1987; Weemstra et al. Reference Weemstra, Peay, Davies, Mohamad, Itoh, Tan and Russo2020), and with slope position in catenary toposequences (Borden et al. Reference Borden, Baillie and Hallett2020).

Soils are complex ecosystems, and host numerous and diverse fauna, the activities of which affect the local distributions and availabilities of abiotic resources. They spatially re-arrange the soil’s solid components and modify pore systems, water retention and release characteristics, and the forms and availabilities of nutrients (Wood, Reference Wood1988; Lee & Wood, Reference Lee and Wood1971; Barros et al. Reference Barros, Cumi, Hallaire, Chauvel and Lavelle2001). The most important and widespread agents of faunal pedoturbation in tropical forests are earthworms (Goodnight & Goodnight,Reference Goodnight and Goodnight1956; Nemeth & Herrera, Reference Nemeth and Herrera1982) and termites (Nye, Reference Nye1955; Gokhale et al. Reference Gokhale, Sarma, Bhattacharyya and Dutta1958; Donovan et al. Reference Donovan, Eggleton, Dubbin, Batchelder and Dibog2001; Roose-Amsaleg et al. Reference Roose-Amsaleg, Mora and Harry2005; Tuma et al. Reference Tuma, Fleiss, Eggleton, Frouz, Klimes, Lewis, Yusah and Fayle2019).

Termites considerably modify local soil patterns; both physically by the excavation of subsoil materials and construction of termitaria, and chemically by the concentration of nutrients through foraging and processing of litter from the surrounding forest. Termitaria are complex structures, and can include subterranean birth chambers for queens, fungal gardens and combs (Yamada et al. Reference Yamada, Inoue, Wiwatwitaya, Ohkuma, Kudo, Abe and Atsuko Sugimoto2005); nurseries; and elaborate vanes and galleries for ventilation and thermoregulation. Above-ground, there are mounds and chimneys to enhance ventilation and also roofed protective walkways for foraging workers. The mounds and walkways are constructed from excavated soil particles and cemented with mixtures of saliva and clay (Lee & Wood, Reference Lee and Wood1971; Jungerius et al. Reference Jungerius, Van den Ancker and Mücher1999; Korb, Reference Korb2003). They are harder than the surrounding soils (Omo-Malaka, Reference Omo-Malaka1977; Jouquet et al. Reference Jouquet, Tessier and Lepage2004), and have different capacities to adsorb, store and shed water (Mando et al. Reference Mando, Stroosnijder and Brussaard1996; Konate et al. Reference Konate, Le Roux, Tessier and Lepage1999; Sarr et al. Reference Sarr, Agbogba, Russell-Smith and Masse2001). Termites forage for litter from the surrounding forest, and termitaria tend to accumulate harvested nutrients, but are not as enriched with C, N and P as earthworm casts (Baillie et al. Reference Baillie, Bunyavejchewin, Kaewfoo, Baker and Hallett2018). They usually have substantially higher pH, and exchangeable and total contents of cationic nutrients, especially Ca, than the surrounding soils (Arshad, Reference Arshad1981; Davies & Baillie, Reference Davies and Baillie1988).

Some savanna termites construct substantial towers with basal diameters of greater than 10 m. Their mounds often carry structurally distinct and floristically diverse vegetation, giving mosaics of dense wooded thickets in matrices of grassland or open woodland (Glover et al. Reference Glover, Trump and Wateridge1964; Goodland, Reference Goodland1965; Dean et al. Reference Dean, Milton and Jeltsch1999; Holdo & McDowell, Reference Holdo and McDowell2004; Loveridge & Moe, Reference Loveridge and Moe2004; Joseph et al. Reference Joseph, Seymour, Cumming, Cumming and Mahlangu2013).Termite mounds in tropical forests are generally smaller than those in savannas, but they can still significantly increase short-range soil heterogeneity (Pendleton, 1941, Reference Pendleton1942; Arshad, Reference Arshad1981; Davies & Baillie, Reference Davies and Baillie1988; Donovan et al. Reference Donovan, Eggleton, Dubbin, Batchelder and Dibog2001; Obi & Ogununle, Reference Obi and Ogunkunle2009).

The long-term forest ecological research plot in Deciduous Dipterocarp Forest at Mae Ping in northern Thailand is located on a colluvial plain of deep dystric red clay loams that is dotted with termitaria, the surface mounds of which range in diameter from 0.1 to 10 m. The larger mounds support moderately dense thickets, which are visually different from the more open surrounding forest. We here examine the extent to which the spatial variations in floristics and structures of the vegetation are associated with termitogenic soil heterogeneity (Beaudrot et al. Reference Beaudrot, Du, Kassim, Rejmánek and Harrison2011). The study attempts to clarify and quantify fortuitously observed phenomena, rather than test pre-ordained hypotheses.


Study site

The study site is the eastern 8 ha half of the 16 ha Mae Ping forest ecology plot, which is located in the Mae Ping National Park, Li district, Lamphun province, Northern Thailand, at 17o 37’ N, 98o 50’ E. The nearest meteorological station is at the Bhumibol Dam (170 15’ N., 980 58’ E), about 10 km to the south (Fig. 1), where the mean annual rainfall is about 1000 mm, with a marked dry season from December to May. Mean monthly temperatures are lowest in November-December, at about 22 – 24o C, and peak at about 30o C towards the end of the dry season in March – April (DNPWP, 2008).

Figure 1. Location of Mae Ping Forest Ecology plot: Dashed line – Mae Ping National Park boundary Shaded – Mae Ping river, Bhumibol Dam, & other lakes; Solid square – Mae Ping Forest Ecology Plot.

The plot is located on the Shan-Thai terrane, which is a crustal segment that rifted off the Australasian part of Gondwanaland, migrated northwards, and sutured onto the Eurasian plate during the Palaeozoic (Rhodes et al. Reference Rhodes, Conejo, Benchawan, Titus and Lawson2005; Hara, et al. Reference Hara, Kunihara, Kuroda, Adachi, Kurita, Wakita, Hisada, Charusiri, Charoentitirat and Chaodumrong2010). The main bedrock under the plot is Devonian-Silurian limestone with subordinate quartzitic metasediments (DMR, 1983). The plot is moderately heterogenous with respect to geology, soils and ecology (Table 1), but our study is located wholly on the colluvial plain of deep, dystric red clay loam that covers the eastern half of the plot (Fig. 2).

Table 1. Land and forest types on Mae Ping 16 ha ecological research plot

Figure 2. Land types, termite mounds, and sampling in the Mae Ping plot Dashed grey line – Termite study area C – Colluvial plain of deep, dystric red loam with dark calcareous clay on termite mounds L – Shallow eutric loam on limestone fins M – Shallow dystric loam on metargillite hillocks Large circle – Mound > 1 m diameter: Filled are sampled; open are unsampled Unsampled smaller mounds: Small circle 0.5 – 1 m diameter; Star < 0.5 m diameter.

Deep augering shows that the red clay loam reaches to below 3 m. The nature of the termitaria indicate that it is underlain by calcareous material, presumably derived from limestone. The soils are therefore bi-sequent, with a lithological discontinuity at several metres depth.

The plain drains southwards in shallow zero-order declivities (Strahler, Reference Strahler1957). Infiltration is slower than that suggested by the red soil colours, and there is some surface runoff. There are ephemeral puddles of standing water during heavy rain and there are scattered patches of green lichen (Supplementary Table S1).

Field methods

We characterised and sampled the vegetation and soils on 15 sets of four 10 × 10 m quadrats in the eastern 8 ha study site. Each set consisted of one quadrat on a large Macrotermes annandalei mound, with the other three at 10 m distance out in surrounding non-mound forest, located beyond the drip rings of the mound vegetation, and so as not to impinge on neighbouring mounds. The layout is constrained by the juxtaposition of the mounds and does not permit the characterisation of distance decay functions (Sileshi & Arshad, Reference Sileshi and Arshad2012). Tree density is higher on the mounds, and three quadrats in the surrounding forest were needed to give comparable numbers of stems in the mound and non-mound samples.

The mounds on the 8 ha study site were mapped by diameter classes of < 0.5 m, 0.5 – 1 m, and > 1 m (Fig. 2). Military caste termites were collected from each mound, preserved in 70% ethanol, and identified according to the Thailand Termite Classification (Sornuwat & Wongkhalung, Reference Soranuwat and Wongkhaluang2004).

Trees with diameter at breast height (dbh = 1.3 m) > 4.5 cm were identified and measured for dbh in the 10 × 10 m quadrats. Saplings (height > 1.3 m, dbh < 4.5 cm) were identified in a 5 m × 5 m sub-quadrat at the centre of each 10 × 10 m quadrat; and seedlings (height < 1.3 m) were identified in four 1 m × 1 m mini-quadrats in each 10 m × 10 m quadrat.

The topsoil (0–15 cm) at the centre of each quadrat was characterised in the field for Munsell colour and hand texture, and tested for free carbonates with HCl. A disturbed topsoil sample from each quadrat was collected for analysis, with the samples from the three non-mound quadrats in each set bulked for analysis. Undisturbed cores of 100 ml volume were taken from 0 to 10 cm depth at the centre of each 10m x 10m quadrat for further physical analyses.

Soil analyses


The disturbed topsoil samples were analysed at the Soil Analysis Laboratory, Department of Soil Science, Kasetsart University, Bangkok, by the following methods: (1) pH electrometrically in a 1:1 suspension of fresh soil in water; (2) organic C by Walkley-Black acid dichromate oxidation; (3) total N by micro-Kjeldahl distillation; (4) available P by Bray and Kurtz extraction and colourimetric assay; (5) exchangeable cations by leaching with 1M neutral ammonium acetate and assay by atomic adsorption spectrometry (AAS); (6) cation exchange capacity (CEC) by displacement of the sorbed NH4 + with excess NaCl, followed by distillation. Total cations and total P were extracted by digestion with concentrated sulphuric acid and a selenium catalyst, and assayed by AAS (cations) or colourimetry (P). (7) Total micronutrients were extracted by digestion with nitric and perchloric acids and assayed by AAS. (8)


Soil particle size distributions in the disturbed samples were determined by hydrometer after organic matter removal with hydrogen peroxide, carbonate removal with HCl, and clay dispersion with sodium hexametaphosphate (Hesse, Reference Hesse1955).

The undisturbed core samples were analysed at the Department of Silviculture, Faculty of Forestry, Kasetsart University, Bangkok, for: (1) moisture content (MC) by loss in core weight after oven drying; (2) dry bulk density (BD) from core weight after drying; (3) particle density (PD) by pycnometer; (4) porosity is derived from the bulk and particle densities (i.e. porosity % = 100 (1 – (BD/PD)); and (5) saturated hydraulic conductivity (Ks).

Data analyses

Each species was characterised separately in each of the three size classes for: (1) relative frequency, as the percentage of quadrats on which the species occurs; (2) relative density, as the number of stems of a species as a percentage of the total stems of all species; and (3) relative basal area of trees, as the species basal area as a percentage of the total basal area of all tree species. A species’ importance value index (IVI) was estimated as the sum of (relative frequency + relative density + relative basal area) (Curtis & McIntosh, Reference Curtis and McIntosh1951). The Sorensen index (W) of floristic similarity between quadrats A and B was estimated from the number of shared species relative to the total number of species, i.e. 2c/(a + b + 2c), where ‘a’ is the number of species found only on quadrat A, ‘b’ is the number of species found only on quadrat B, and ‘c’ is the number of species common to both (Sorensen, Reference Sorensen1948). Species diversity was characterised by the Shannon-Wiener index, H (= ∑ [(pi × ln(pi)] where pi is the relative density for each species). Floristic cluster analysis of quadrats was based on the Sorenson similarity index (Kent & Coker, Reference Kent and Coker1994). Canonical Correspondence Analysis (CCA) segregated quadrats on floristic composition and also clarified environmental relationships (McCune & Mefford, Reference McCune and Mefford1999).

The laboratory analyses of the mound and non-mound topsoils were compared by ‘t tests’. The associations between quadrat floristics and topsoil properties were tested by Pearson and rank correlations of the soil variables with the main floristic CCA axis for each size class(Palmer, Reference Palmer1993; ter Braak & Verdonschot, 1993; Uurito et al. Reference Uurtio, Monteiro, Kandola, Shawe-Taylor, Fernandez-Reyes and Rousu2017). The stoichiometries of the exchangeable cations and pH were compared visually in Alvim (Reference Alvim1978) roses.


Termite mounds

We mapped and measured 89 termite mounds in the study area (Fig. 2), giving an approximate mean density of 11/ha (Table 2). This is equivalent to a mean inter-mound spacing of about 30 m. The actual spacing is variable and Fig. 2 shows that distances between adjacent large mounds range from 20 to 70 m. About three quarters (64) of the mounds were of intermediate size (diameter 0.5 – 1 m). Soldiers of the fungivorous species Macrotermes annandalei (Macrotermitinae) (Davies, Reference Davies1997; Hyodo et al. Reference Hyodo, Tayasu, Inoue, Azuma, Kudo and Abe2003) were found in 81 mounds, including all 18 in the largest diameter class (>1 m).

Table 2. Termites in mounds on Mae Ping 8 ha study site

Soldiers of Globitermes sulphureus (Amitermitinae) (Yamada et al. 2003, 2004) were found in four mounds of intermediate size. No termites were observed in four apparently abandoned mounds of intermediate size (Table 2).

The mounds of Macrotermes annandalei account for about 96% of the total soil volume excavated (Tuma et al. Reference Tuma, Fleiss, Eggleton, Frouz, Klimes, Lewis, Yusah and Fayle2019). They have irregular profiles, with diameters of up to 10 m and heights of up to 2.5 m (Fig. 3). The few Globitermes sulphureus mounds are roughly cylindrical and do not attain great size, ranging in diameter and height from 40 cm to 1 m They are dome-topped and have near-vertical sides, which are not readily colonised by plants. They are formed of red clay loam that appears to have been excavated from the upper subsoil (Supplementary Figure S1).

Table 3. Forest structure at Mae Ping

Where Dbh = diameter at breast height (1.3 m).

Figure 3. Macrotermes annandalei mound in Deciduous Dipterocarp Forest, Mae Ping.


The density of large trees in the non-mound forest is moderate (Table 3). The canopy, at heights of 20 – 30 m, is almost closed in the wet season, but there is sufficient light at ground level to allow some cover of grasses, forbs, and a few sedges. The mound vegetation forms lower and denser thickets with a trebling of sapling density compared to the non-mound forest and an increase in the ratio of small: large trees from 0.56 to 1.14 (Table 3 and Fig. 3). The structural differences are paralleled in the floristic composition. The commonest tree species in the non-mound forest is Dipterocarpus tuberculatus, with Shorea obtusa subordinate. Frequent non-dipterocarp trees include Aporosa villosa, Quercus kerrii, Strychnos nux-blanda, Buchanania lanzan, and Symplocos racemosa. The composition of the seedlings is similar, and is dominated by the same dipterocarps, but there is a greater range of non-dipterocarps (Table 4a). The intermediate sapling class contains some dipterocarps, similar to those in the trees, but is dominated by the non-dipterocarps Anacolosa ilicoides and Dillenia obovata.

Table 4a. Main species in quadrats in non-mound forest, Mae Ping

The floristic composition of the vegetation on the termite mounds differs distinctly from the surrounding forest, for all size classes. There are few dipterocarps, and the common tree species include Morinda coreia, Ziziphus cambodiana, Schleichera oleosa, Walsura villosa, and Siphonodon celastrineus (Table 4b; Supplementary Table S2b).

Table 4b. Main species in quadrats on termite mounds, Mae Ping

The diversity and similarity indices (Table 5) confirm the floristic differences between the mound and non-mound vegetation, with few shared species and little similarity between the groups, especially for the trees. The differences are also clearest for the trees in the cluster analyses (Fig. 4a–4c). There are two main clusters for each of the non-mound and mound quadrats. The dipterocarps Dipterocarpus tuberculatus and Shorea obtusa dominate in both of the non-mound clusters, and these are differentiated on the non-dipterocarps. The commonest non-dipterocarps in the quadrats of cluster 1 are Aporosa villosa, Quercus kerrii and Strychnos nux-blanda, whereas Symplocos racemosa and Dalbergia dongnaiensis are important in cluster 2.

Table 5. Floristic diversity and similarity, Mae Ping

Figure 4. Floristic cluster analyses of quadrats at Mae Ping (a) Trees (b) Saplings (c) Seedlings.

Frequent tree species in the quadrats of cluster 1 for the mound quadrats include Morinda coreia, Antidesma ghaesembilla, Ziziphus cambodiana, Cassia fistula, and Grewia eriocarpa., whereas Schleichera oleosa, Walsura villosa, Siphonodon celastrineus, and Xantolis burmanica are common trees in mound cluster 2. The separation between the non-mound and mound quadrats is complete for the seedlings but is less pronounced than for the larger size classes (Fig. 4c). The non-mound saplings are dipterocarp-poor and more diverse than the other size classes, but the sapling analysis still separates the two formations, with four clusters for the mound and five for the non-mound quadrats (Fig. 4b).

The Canonical Correspondence Analyses (CCA) confirms the clear floristic separation of mound and non-mound vegetation. The CCA patterns in the non-mound quadrats are similar for the trees and seedlings, with tight clustering on both axes (Fig. 5a & 5c). The wider spread for the saplings on both axes confirms that this size class is more diverse (Fig. 5b). The greater diversity of the mound vegetation gives wider spread on both axes.

Figure 5. Canonical Correspondence Analyses (CCA) with soil correlations at Mae Ping (a) Trees (b) Saplings (c) Seedlings Open rectangle – Quadrat in non-mound forest; Filled triangle – Quadrat on termite mound; Lines – Pearson correlations of topsoil variable with CCA axis 1.


The non-mound topsoils are brownish red clay loams with compound blocky structures breaking to crumb (Supplementary Table S1). The subsoils are bright red, friable light clays, at least 3.5 m deep. There are no mottles in the subsoils, but convoluted layers of brittle black manganiferous angular concretions occur at several depths. These soils are moderately acid and dystric, with base saturations in the topsoils of almost 50%, but the subsoils are more acid, with base saturations of less than 35%. All of the non-mound soils are inert to HCl and contain no free carbonates.

The mound soils are darker, with yellower Munsell hues, and lower chromas and values. Some contain enough free carbonates to effervesce slightly with HCl. They are clay textured throughout, crack more readily, and have blockier structures than the non-mound soils. Their subsoils are also darker and yellower, with predominant Munsell hues of 5YR to 10YR compared with 10R and 2.5 YR for the non-mound soils. However, organic darkening gives both groups overlapping ranges of brownish topsoil colours.

Chemically, the non-mound and mound topsoils differ little with respect to organic C and both available and total P (Table 6). The mound topsoils contain significantly more total N, but the difference is not substantial. However, the two sets differ substantially and significantly with respect to base status. pH values are at least a unit (i.e. ten-fold) higher in the mound topsoils, and this is accompanied by a highly significant doubling of base saturation. All of the main exchangeable cations are very significantly higher in the mounds, with K and Ca approximately quadrupled, and Mg and CEC doubled (Table 6). The stoichiometries of exchangeable cations are depicted in roses (Fig. 6). These are based on the idea of wind roses, and show nutrient proportions as well as quantities (Alvim, Reference Alvim1978). Comparison of the sizes of the polygons confirms the substantially larger contents of exchangeable cations in the mound soils. The two polygons are of different shape, the comparison of which shows how the proportions of cations vary between the sets. Enrichment of the mound soils with Ca widens the Ca:Mg ratio from 2 to 4.5 (Table 6), but this is not thought to be sufficient to induce Mg deficiency (Osemwota et al. Reference Osemwota, Omueti and Ogboghodo2007; Baillie et al. Reference Baillie, Bunyavejchewin, Kaewfoo, Baker and Hallett2018).

Table 6. Comparison of non-mound and mound topsoils

*** p < 0.001;

** p < 0.01;

* p < 0.05; ns p > 0.05.

Figure 6. Stoichiometric roses of exchangeable cations and pH in Mae Ping topsoils Shaded inner: Non-mound topsoil Clear outer: Mound topsoil All scales, except pH, are linear.

Some cations also show marked differences in their total, as well as exchangeable, contents (Table 6). Total Mg and Ca are significantly higher in the mound soils. Mean total Fe is substantially higher in the mound soils, but the high variability of Fe in the non-mound soils renders the difference non-significant. Totals of the micro-nutrients Zn and Cu are significantly higher in the mound soils, but total Mn contents are almost identical.

The laboratory soil particle size distributions confirm the field textures, with significantly higher clay and lower sand in the mound topsoils. The mound topsoils have significantly lower mean particle densities, but significantly higher bulk densities. This results in significantly lower overall porosities and contributes to significantly lower saturated hydraulic conductivities (Table 6).

Soil – forest associations

The differences between the termite mounds and the surrounding forest are visually clear for both vegetation and soils. The associations between soils and vegetation are confirmed by the soil ‘ t tests’ (Table 6) and by the canonical correspondence (CCA) analyses (Fig. 5a–5c). The soil base status variables are correlated with CCA Axis 1 for all three size classes (Table 7). Total manganese differs in that it is aligned subparallel to Axis 2 in the tree CCA (Fig. 5a). This accords with the almost identical mean total Mn values for the two groups and the non-significant ‘t test’ for Mn (Table 6).

Table 7. Correlations of topsoil variables against CCA axes 1 and 2 for different size classes



The non-mound forest is Deciduous Dipterocarp Forest (DDF), a formation that is adapted to seasonal moisture stress and frequent fires. It is widespread in the drier parts of northern Thailand, Upper Myanmar and northern Kampuchea (Bunyavejchewin et al. Reference Bunyavejchewin, Baker, Davies, McShea, Davies and Bhumpakphan2011; Wohlfahrt et al. Reference Wohlfart, Wegmann and Leimgruber2014; Nguyen & Baker Reference Nguyen and Baker2016). Its seasonally open canopy and significant ground cover provide some grazing for large herbivores, such as elephants and ungulates. The main dipterocarps in the 20–30 m tall canopy are Dipterocarpus obtusifolius, D. tuberculatus, Shorea siamensis and S. obtusa, the proportions of which vary locally (Table 3 & Supplementary Table S2a; Fig. 4a). There are three DDF subtypes within the 16-hectare plot at Mae Ping, on different bedrock and soils (Table 1 and Fig. 2). The scarcity of dipterocarp saplings in the non-mound quadrats on the 8 ha study site may be due to a poor supply of seeds in the recent past, as the dipterocarps flower and fruit irregularly and gregariously. Prolonged droughts and the scarcity of bare sites suitable for germination may also be involved (Nguyen & Baker, Reference Nguyen and Baker2016).

The mound vegetation is more typical of Mixed Deciduous Forest (MDF), a fire-adapted formation with few or no dipterocarps that is often associated with calcareous lithologies and is widespread in seasonal climates in northern Thailand and adjacent countries (Marod et al. Reference Marod, Kutintara, Yarwudhi, Tanaka and Nakashisuka1999; Bunyavejchewin et al. 2011). Neither of the canopy species that normally differentiate the main MDF sub-types, i.e. Tectona grandis and Lagerstroemia calyculata, (Bunyavejchewin, Reference Bunyavejchewin1983), were recorded on the mound quadrats. This may be because the stands are only as old as the mounds, and have yet to acquire the canopy composition f mature MDF. However, the floristic correspondence of the mound vegetation with lower canopy and ground layers of typical MDF is clear.


The contrast between the two sets of soils is enhanced by the lithological discontinuity in the regolith. The non-mound soils are developed in deep dystric colluvium several metres deep, whereas the larger mounds include calcareous materials that have been excavated from the limestone contact. The lithological difference means that the two groups of soils develop along separate pedogenic trajectories and qualify for different high-level taxa in the international soil classification systems. The non-mound soils are Acric Ferralsols in the World Reference Base (IUSS, 2022) and Rhodic Kandiudoxes in the USDA Soil Taxonomy (Soil Survey Staff, 2022). The mound soils are Provertic Cambisols in the World Reference Base and Vertic Eutrodepts in Soil Taxonomy.

Clay contents increase with depth in the non-mound soils (Supplementary Table S1), although mostly not enough to qualify the subsoils as argic or argillic horizons sensu stricto. The finer textures of the mound topsoils could just be due to indiscriminate excavation of clay-enriched subsoil material. However, termite selectivity for particle size may contribute to the higher clay contents in the mound topsoils, as many termites are selective in their choice of particles for excavation and construction, with coarse sand particles carried by their mandibles and finer in their crops (Stoops, Reference Stoops and Bouillon1964; Abe, et al. Reference Abe, Yamamoto and Wakatsuki2009). The lower total porosities and hydraulic conductivities of the mound soils are unexpected, because the ventilation macro-voids should increase overall porosity. Micromorphological analysis might show that the low values are due to dense packing in the construction process. They contrast with findings that overall porosity increases in savanna termitaria (e.g. Konate et al. Reference Konate, Le Roux, Tessier and Lepage1999). The consequently low infiltration and high surface runoff of the Mae Ping mounds are consistent with the lichen mats and morphological indicators of surface puddling.

Hard rounded gravels of concretionary manganese dioxide (MnO2) are common in tropical forest subsoils. The red non-mound clay loams at Mae Ping have abnormally high contents of free MnO2, distributed in convoluted layers of concretions that are unusually angular and brittle (Supplementary Table S1). MnO2 concretions are often associated with fluctuating soil aeration, but the matrices of the non-mound subsoils are bright red and unmottled, and the profiles appear to be freely drained. Similar concretions have been seen in unmottled red clays over limestone in other tropical biomes, such as forest in Belize (Baillie et al. Reference Baillie, Wright, Holder and Fitzpatrick1993), savanna in Tanzania (AHT, 1980), and semi-desert in Jordan (HTS, 1994), suggesting that a calcareous input into the soil parent material, and consequently elevated pH, base and Ca status, may be implicated in their formation.

Total Mn is unique among the cations assayed at Mae Ping, in that there is no difference observed between the calcareous mound and non-calcareous non-mound topsoils (Table 2). Also, total Mn is weakly correlated with CCA Axis 2, unlike the basic cations which are correlated with Axis 1 (Fig. 5a). The effect of soil Mn on the forest at Mae Ping is unclear. Manganese has been identified as a possible micronutrient but it can be toxic at high concentrations. It has been associated with P mobilisation and uptake, and can have complex interactions with cationic nutrients, especially Ca (Le Mare, Reference Le Mare1977; Hall & Huang Reference Hall and Huang2017; Zemunik et al. Reference Zemunik, Winter and Turner2020).

The termites forage and import surface litter from surrounding areas for consumption by their symbiotic fungi. The consequently intense metabolic activity has been reported to increase subsoil temperatures, and the partial pressures of CO2, methane and other respiratory products in the soil atmosphere (Seiler et al. Reference Seiler, Conrad and Scharffe1984).

Forest – soil associations

The distributions of the contrasting DDF and MDF-like thickets in the study area are clearly associated with the soil differences. As in many tropical forests, it is not possible to determine definitively which soil attributes most influence forest pattern, because many are highly inter-correlated. Soil processes and attributes that might be affected by the higher pH, bases and Ca in the mound soils include: lability of Al; stimulation of microbial activity; interactions with Mn; stoichiometric suppression of K and Mg by copious Ca uptake; and possibly sequestration or solubilisation of P (Baillie et al. Reference Baillie, Bunyavejchewin, Kaewfoo, Baker and Hallett2018). Unusually for tropical forests, both available and total phosphorus are insignificant as differentiae at Mae Ping (Vitousek & Sanford, Reference Vitousek and Sanford1986).

Pedoturbation and forest diversification

The lithological discontinuity and the excavation of distinctly calcareous regolith material enhances the contrast in base status between the mound and non-mound soils. However, this trend is also apparent in tropical forest termite mounds derived from lithologically uniform regoliths (Davies & Baillie, Reference Davies and Baillie1988), because weathering rock C horizons often have higher base status than overlying leached subsoil B horizons (Baillie, Reference Baillie and Richards1996).

The termitogenic importation of substrate cations into the mound topsoils increases the amounts available for active biological cycling. This counters the climatogenic trend towards topsoil dystrophication by downward leaching of soil bases. Upward translocation by termite excavation can modify or retard other leaching-driven pedological processes, such as argilluviation, mineral weathering and desilication (Baillie, Reference Baillie and Richards1996).

Although widespread and important, termites are not the only agents of pedoturbation that renew soil diversity and counter leaching-driven dystrophication in tropical forests. Excavation and modification of forest soils by earthworms has been noted in forests on skarn rocks in western Thailand (Pendleton, 1941, Reference Pendleton1942; Goodnight & Goodnight,Reference Goodnight and Goodnight1956; Nemeth & Herrera, Reference Nemeth and Herrera1982; Baillie et al. Reference Baillie, Bunyavejchewin, Kaewfoo, Baker and Hallett2018; Van Groenigen et al. Reference Van Groenigen, Van Groenigen, Koopmans, Stokkermans, Vos and Lubbers2019). Other pedoturbation agents include: pangolins (Shrestra et al. Reference Shrestha, Bhattarai, Shrestha and Koju2021), megapodes (Machincote et al. Reference Machincote, Brauch and Villareal2004; Karawita et al. Reference Karawita, Perera, Gunawardane and Dayawansa2018), wasps, cicadas (Tuma et al. Reference Tuma, Fleiss, Eggleton, Frouz, Klimes, Lewis, Yusah and Fayle2019), mud lobsters (Hossain et al. Reference Hossain, Bujang, Kamal, Zakaria, Muslim and Nadzri2019); ants (Green et al. Reference Green, Pettry and Switzer1999); and wild boar (Ickes et al. Reference Ickes, Paciorek and Thomas2005). Landslides and treefall also excavate soils and chemically replenish mineral nutrients by exposing fresh regolith to weathering (Heineman et al. Reference Heineman, Russo, Baillie, Mamit, Chai, Chai, Hindley, Lau, Tan and Ashton2015).


Our results indicate that the floors of some dry tropical forests are mosaics of stable dystric soils interspersed with patches that have been rejuvenated by termite pedoturbation. This pattern somewhat mirrors the spatial mosaic of mature canopy and gaps in the forest canopy above. Termite pedoturbation also occurs in moist and evergreen forests (Baillie, Reference Baillie and Richards1996). The rejuvenation alters the edaphic conditions for the forest with respect to soil structure, root aeration, moisture dynamics, quantities and availability of nutrients, and mechanical stability for the root environment. At Mae Ping these changes are sufficient to significantly influence the local structural and floristic diversity of the forest. Pedoturbation and soil diversity, both as an instantaneous condition and as a set of ongoing processes, need to be accommodated at a range of spatial scales in comprehensive paradigms of tropical forest dynamics.

Supplementary material

To view supplementary material for this article, please visit

Data availability statement

The data can be downloaded from


We are grateful for constructive comments from Brian Kerr, Wayne Borden, Professor Peter Ashton, and the journal’s anonymous reviewers. We acknowledge use of the World Soil Survey Archive and Catalogue (WOSSAC), Cranfield University, UK. MK and IB received logistic support from the Royal Forest Department, Thailand; Harvard Forest; and from the former Center for Tropical Forest Research (now ForestGEO), USA.

Author contributions

MK did the detailed fieldwork and data analyses, commissioned the soil analyses, wrote the Thai text, and drafted the original graphics. SB, DM and DW supervised the survey design, data analyses and Thai text. IB, MK, SB, SH and SD wrote and modified the English text and graphics and undertook data management.

Financial support

The work undertaken as members of National Parks Wildlife and Plant Conservation Department of the Royal Government of Thailand (MS and SB); Bullard Fellowship from Harvard Forest, and a travel grant from Centre for Tropical Forrest Studies (IB).

Competing interests

There is no conflict of interest associated with this publication.


Abe, S, Yamamoto, S and Wakatsuki, T (2009) Soil-particle selection by the mound-building termite Macrotermes bellicosus on a sandy loam soil catena in a Nigerian tropical savanna. Journal of Tropical Ecology 25, 449452. Scholar
AHT (1980) Land suitability survey. Phase I. Annexes 2 & 3. National Coconut Development Programme, Tanzania. Agrar-und-Hydrotechnik GmbH, Essen. Available at Scholar
Alvim, PdeT (1978) Perspectiva de producção agricola na região Amazonica. Interçiencia 3, 343349.Google Scholar
Arshad, MA (1981) Physical and chemical properties of termite mounds of two species of Macrotemes (Isoptera, Termitidae) and the surrounding soils of the semi-arid savanna of Kenya. Soil Science 132, 161174.CrossRefGoogle Scholar
Baillie, IC (1996) Soils of the humid tropics. In Richards, PW (ed), The Tropical Rain Forest: An Ecological Study, 2nd Edn. Cambridge: Cambridge University Press, pp. 256286.Google Scholar
Baillie, IC, Ashton, PS, Court, MN, Anderson, JAR, Fitzpatrick, EA and Tinsley, J (1987) Site characteristics and the distribution of tree species in Mixed Dipterocarp Forest on Tertiary sediments in central Sarawak, Malaysia. Journal of Tropical Ecology 3, 201220. CrossRefGoogle Scholar
Baillie, IC, Bunyavejchewin, S, Kaewfoo, M, Baker, PJ and Hallett, SH (2018) Stoichiometry of cationic nutrients in Phaeozems derived from skarn and Acrisols from other parent materials in lowland forests of Thailand. Geoderma Regional 12, 19. CrossRefGoogle Scholar
Baillie, IC, Wright, ACS, Holder, MA and Fitzpatrick, EA (1993) Revised classification of the soils of Belize. Bulletin 49. Natural Resources Institute, Chatham. Available at Google Scholar
Barros, E, Cumi, P, Hallaire, V, Chauvel, A and Lavelle, P (2001) The role of macrofauna in the transformation and reversibility of soil structure of an oxisol in the process of forest to pasture conversion. Geoderma 100, 193213.CrossRefGoogle Scholar
Beaudrot, L, Du, Y-J., Kassim, AR, Rejmánek, M and Harrison, RD (2011) Do epigeal termite mounds increase the diversity of plant habitats in a tropical rain Forest in Peninsular Malaysia? PLoS One 6, 5,, (e19777).CrossRefGoogle Scholar
Borden, RW, Baillie, IC and Hallett, SH (2020) The East African contribution to the formalisation of the soil catena concept. Catena 185, 104291. CrossRefGoogle Scholar
Bunyavejchewin, S (1983) Analysis of tropical dry deciduous forest of Thailand. I. Characteristics of dominance types. Natural History Bulletin Siam Society 31, 109122.Google Scholar
Bunyavejchewin, S., Baker, PJ and Davies, SJ (2011) Seasonally dry tropical forests in continental Southeast Asia: structure, composition and dynamics. In McShea, WJ, Davies, SJ and Bhumpakphan, N (eds), The Ecology and Conservation of Seasonally Dry Forests in Asia. Washington, DC: Smithsonian Press, pp. 935.Google Scholar
Curtis, JT and McIntosh, RP (1951) An upland forest continuum in the prairie-forest border region of Wisconsin. Ecology 32, 476496.CrossRefGoogle Scholar
Davies, AG and Baillie, IC. (1988) Soil-eating by red leaf monkey (Presbytis rubicunda) in Sabah, northern Borneo. Biotropica 20, 252258. CrossRefGoogle Scholar
Davies, RG (1997) Termite species richness in fire-prone and fire-protected dry deciduous Dipterocarp Forest in Doi Suthep-Pui National Park, northern Thailand. Journal Tropical Ecology 13, 153160. CrossRefGoogle Scholar
Dean, WRJ, Milton, SJ and Jeltsch, F (1999) Large trees, fertile islands, and birds in arid savanna. Journal Arid Environments 41, 6178. CrossRefGoogle Scholar
DMR (1983) Geological map of Thailand. 1:500,000. Northern Sheet. Bangkok: Department of Mineral Resources.Google Scholar
Donovan, SE, Eggleton, P, Dubbin, WE, Batchelder, M and Dibog, L (2001) The effect of a soil-feeding termite, Cubitermes fungifaber (Isoptera: Termitidae) on soil properties: termites may be an important source of soil microhabitat heterogeneity in tropical forests. Pedobiologia 45, 111.CrossRefGoogle Scholar
DNPWP (2008) 30-Year Climate Statistics. Bangkok: Department of National Parks, Wildlife and Plants.Google Scholar
Glover, PE, Trump, EC and Wateridge, LED (1964) Termitaria and vegetation patterns on the Loita plains of Kenya. Journal of Ecology 52, 365377.CrossRefGoogle Scholar
Gokhale, NG, Sarma, SN, Bhattacharyya, NG and Dutta, JS (1958) Effect of termite activity on the chemical properties of tea soils. Scientific Culture 24, 229.Google Scholar
Goodland, RJA (1965) On termitaria in a savanna ecosystem. Canadian Journal Zoology 43, 641650.CrossRefGoogle Scholar
Goodnight, CL and Goodnight, ML (1956) Some observations on tropical rainforest in Chiapas, Mexico. Ecology 37, 139150.CrossRefGoogle Scholar
Green, WP, Pettry, DE and Switzer, RE (1999) Structure and hydrology of mounds of imported fire ants in the south-eastern United States. Geoderma 93, 117. CrossRefGoogle Scholar
Hall, SJ and Huang, W (2017) Iron reduction: a mechanism for dynamic cycling of occluded cations in tropical forest soils? Biogeochemistry 136, 91102. CrossRefGoogle Scholar
Hara, H, Kunihara, T, Kuroda, J, Adachi, Y, Kurita, H, Wakita, K, Hisada, K, Charusiri, P, Charoentitirat, T and Chaodumrong, P (2010) Geological and geochemical aspects of a Devonian siliceous succession in northern Thailand: implications for the opening of the Paleo-Tethys. Palaeogeography, Palaeoclimatology, Palaeoecology 297, 452464. CrossRefGoogle Scholar
Heineman, KD, Russo, SE, Baillie, IC, Mamit, JD, Chai, PPK, Chai, L, Hindley, EW, Lau, BT, Tan, S and Ashton, PS (2015) Evaluation of stem rot in 339 Bornean tree species: implications of size, taxonomy, and soil-related variation for aboveground biomass estimate. Biogeosciences 12, 57355751. CrossRefGoogle Scholar
Hesse, PR (1955) A chemical and physical study of the soils of termite mounds in East Africa. Journal of Ecology 43, 449461.CrossRefGoogle Scholar
Holdo, RM and McDowell, LR (2004) Termite mounds as nutrient rich food patches for elephants. Biotropica 36, 231239. Google Scholar
Hossain, MS, Bujang, JS, Kamal, AHM, Zakaria, MH, Muslim, AM and Nadzri, MI (2019) Effects of burrowing mud lobsters (Thalassina anomala Herbst 1804) on soil macro- and micronutrients in a Malaysian mangrove. Estuarine, Coastal and Shelf Science 228. CrossRefGoogle Scholar
HTS (1994) The soils of Jordan. Level 2. Semi-detailed studies. Hemel Hempstead, UK: Hunting Technical Services.Google Scholar
Hyodo, F, Tayasu, I, Inoue, T, Azuma, J-I, Kudo, T and Abe, T (2003) Differential role of symbiotic fungi in lignin degradation and food provision for fungus-growing termites (Macrotermitinae: Isoptera). Functional Ecology 17, 186193. CrossRefGoogle Scholar
Ickes, K, Paciorek, CJ and Thomas, SC (2005) Impacts of nest construction by native pigs (Sus scrofa) on lowland Malaysian rain forest saplings. Ecology 86, 15401547. CrossRefGoogle Scholar
IUSS Working Group WRB (2022) World Reference Base for Soil Resources, 4th Edn. Vienna: International Union of Soil Sciences.Google Scholar
Joseph, GS, Seymour, CL, Cumming, GS, Cumming, DHM and Mahlangu, Z (2013) Termite mounds as islands: woody plant assemblages relative to termitarium size and soil properties. Journal of Vegetation Science 24, 702711. https://doi:10.1111/j.1654-1103.2012.01489.x CrossRefGoogle Scholar
Jouquet, P, Tessier, D and Lepage, M (2004) The soil structural stability of termite nest: role of clays in Macrotermes bellicosus (Isoptera, Macrotermitinae) mound soils. European Journal of Soil Biology 40, 2329. CrossRefGoogle Scholar
Jungerius, PD, Van den Ancker, JA and Mücher, HJ. (1999) The contribution of termites to the microgranular structure of soils on the Uasin Gishu Plateau, Kenya. Catena 34 349363.CrossRefGoogle Scholar
Karawita, H, Perera, P, Gunawardane, P and Dayawansa, N (2018) Habitat preference and den characterization of Indian Pangolin (Manis crassicaudata) in a tropical lowland forested landscape of southwest Sri Lanka. PLOS 13. Scholar
Kent, M and Coker, P (1994) Vegetation Description and Analysis. New York: John Wiley & Sons Ltd. Google Scholar
Konate, SX, Le Roux, D, Tessier, D and Lepage, M (1999) Influence of large termitaria on soil characteristics, soil water regime and tree leaf shedding in a West African savanna. Plant and Soil 206, 4760. CrossRefGoogle Scholar
Korb, J (2003) Thermoregulation and ventilation of termite mounds. Naturwissenschaften 90, 212219. https://doi.10.1007/s00114-002-0401-4 CrossRefGoogle ScholarPubMed
Lee, KE and Wood, TG (1971) Termites and Soils. London and New York: Academic Press.Google Scholar
Le Mare, PH (1977) Experiments on effects of phosphorus on the manganese nutrition of plants. III. The effect of calcium:phosphorus ratio on manganese in cotton grown in Buganda soil. Plant and Soil 47, 621630.CrossRefGoogle Scholar
Loveridge, JP and Moe, SR (2004) Termitaria as browsing hotspots for African megaherbivores in miombo woodland. Journal Tropical Ecology 20, 337343. CrossRefGoogle Scholar
Machincote, M, Brauch, LC and Villareal, D (2004) Burrowing owls and burrowing mammals are ecosystem engineers interchangeable as facilitators. Oikos 106, 527535. CrossRefGoogle Scholar
Mando, A, Stroosnijder, L and Brussaard, L (1996) Effect of termites on infiltration into crusted soil. Geoderma 74, 107113. CrossRefGoogle Scholar
Marod, D, Kutintara, U, Yarwudhi, C, Tanaka, H and Nakashisuka, T (1999) Structural dynamics of a natural mixed deciduous forest in western Thailand. Journal of Vegetation Science 10, 777786. CrossRefGoogle Scholar
McCune, B and Mefford, MJ (1999) PC – ORD Multivariate Analysis of Ecological Data: Version 4 for Windows. Gleneden Beach, Oregon: MjM Software Design.Google Scholar
Nemeth, A and Herrera, R (1982) Earthworm populations in a Venezuelan tropical rainforest. Pedobiologia 23, 437443. Available at Scholar
Nguyen, TT and Baker, PJ (2016) Structure and composition of deciduous Dipterocarp Forest in Central Vietnam: patterns of species dominance and regeneration failure. Plant Ecology & Diversity 9, 589601. CrossRefGoogle Scholar
Nye, PH (1955) Some soil-forming processes in the humid tropics. IV. The action of the soil fauna. Journal of Soil Science 6, 7383. CrossRefGoogle Scholar
Obi, JL and Ogunkunle, AO (2009) Influence of termite infestation on the spatial variability of soil properties in the Guinea savanna region of Nigeria. Geoderma 148, 357363. Scholar
Omo-Malaka, SL (1977) A note on the bulk density of termite mounds. Australian Journal of Soil Research 15, 9394. CrossRefGoogle Scholar
Osemwota, IO, Omueti, JAI and Ogboghodo, AI (2007) Effect of calcium/magnesium ratio in soil on magnesium availability, yield, and yield components of Maize. Communications in Soil Science and Plant Analysis 38, 1920, 2849–2860. CrossRefGoogle Scholar
Palmer, MW (1993) Putting things in even better order: the advantages of canonical 5 correspondence analysis. Ecology 74, 22152230. CrossRefGoogle Scholar
Pendleton, RL (1941) Some results of termite activity in Thailand soils. Thailand Science Bulletin 3, 2953.Google Scholar
Pendleton, RL (1942) Importance of termites in modifying certain Thailand soils. Journal of the American Society of Agronomy 34, 340344. CrossRefGoogle Scholar
Rhodes, BP, Conejo, R, Benchawan, T, Titus, S and Lawson, R (2005) Palaeocurrents and provenance of the Mae Rim Formation, Northern Thailand: implications for tectonic evolution of the Chiang Mai basin. Journal of the Geological Society 162, 5163. CrossRefGoogle Scholar
Richards, PW (1996) The Tropical Rain Forest: An Ecological Study, 2nd Edn. Cambridge: Cambridge. University Press.Google Scholar
Roose-Amsaleg, C, Mora, P and Harry, M (2005) Physical, chemical and phosphatase activities characteristics in soil-feeding termite nests and tropical rainforest soils. Soil Biology & Biochemistry 37, 19101917. CrossRefGoogle Scholar
Sarr, MC, Agbogba, A, Russell-Smith, A and Masse, D (2001) Effect of soil faunal activity and woody shrubs on water infiltration rates in a semi-arid fallow of Senegal. Applied Soil Ecology 16, 283290.CrossRefGoogle Scholar
Seiler, W, Conrad, R and Scharffe, D (1984) Field studies of methane emission from termite nests into the atmosphere and measurements of methane uptake by tropical soils. Journal of Atmospheric Chemistry 1, 171186. CrossRefGoogle Scholar
Sileshi, GW and Arshad, MA (2012) Application of distance-decay models for inferences about termite mound-induced patterns in Dryland ecosystems Journal of Arid Environments 77, 138148. CrossRefGoogle Scholar
Shrestha, S, Bhattarai, S, Shrestha, B and Koju, NP (2021) Factors influencing the habitat choice of pangolins (Manis spp.) in low land of Nepal. Ecology and Evolution 11, 1468914696. CrossRefGoogle ScholarPubMed
Soil Survey Staff (2022) Keys to Soil Taxonomy, 13th Edn. Lincoln, Nebraska: USDA-Natural Resources Conservation Service.Google Scholar
Soranuwat, Y and Wongkhaluang, J (2004) Manual for Termite Classification in Thailand. Bangkok: Department of Forestry.Google Scholar
Sorensen, T (1948) A method of establishing groups of equal amplitude in plant sociology based on similarity of species content. Kongelige Danske Videnskabernes Selskabs Skrifter 5, 134.Google Scholar
Stoops, G (1964) Application of some pedological methods to the analysis of termite mounds. In Bouillon, A (ed), Etudes sur les Termites Africains. Leopoldville: Leopoldville University, pp. 379398.Google Scholar
Strahler, AN (1957) Quantitative analysis of watershed geomorphology. Transactions American Geophysical Union 38, 913920.Google Scholar
ter Braak, JF and Verdonschot, PFM (1995) Canonical correspondence analysis and related multivariate methods in aquatic ecology. Aquatic Sciences 57, 255289. CrossRefGoogle Scholar
Tuma, J, Fleiss, S, Eggleton, P, Frouz, J, Klimes, P, Lewis, OT, Yusah, M and Fayle, TM (2019) Logging of rainforest and conversion to oil palm reduces bioturbator diversity but not levels of bioturbation. Applied Soil Ecology 144, 123133. CrossRefGoogle Scholar
Uurtio, V, Monteiro, JM, Kandola, J, Shawe-Taylor, J, Fernandez-Reyes, D and Rousu, J (2017) A Tutorial on canonical correlation methods. Article 95, ACM Computing Surveys, 50.Google Scholar
Van Groenigen, JW, Van Groenigen, KJ, Koopmans, GF, Stokkermans, L, Vos, HMJ and Lubbers, IM (2019) How fertile are earthworm casts? A meta-analysis. Geoderma 338, 525535. CrossRefGoogle Scholar
Vitousek, PM and Sanford, RL Jr (1986) Nutrient cycling in moist tropical forest. Annual Review of Ecology & Systematics 17, 137147.CrossRefGoogle Scholar
Weemstra, M, Peay, KG, Davies, SJ, Mohamad, M, Itoh, A, Tan, S and Russo, SE (2020) Lithological constraints on resource economies shape the mycorrhizal composition of a Bornean rain forest. New Phytologist 228, 253268. CrossRefGoogle ScholarPubMed
Wohlfart, C, Wegmann, M and Leimgruber, P (2014) Mapping threatened dry deciduous Dipterocarp Forest in South-east Asia for conservation management. Tropical Conservation Science 7, 597613.CrossRefGoogle Scholar
Wood, TG (1988) Termites and the soil environment. Biology & Fertility of Soils 6, 228236. CrossRefGoogle Scholar
Yamada, A, Inoue, T, Sugumoto, A, Takematsu, Y, Kumai, F, Fujita, A, Tayasu, T, Klanfkaew, C, Kiributr, N, Kudo, T and Abe, T (2003 & 2004) Abundance and biomass of termites (Insecta:Isoptera) in dead wood in Dry Evergreen Forest in Thailand. Sociobiology 42, 569582 & 43, 231–232.Google Scholar
Yamada, A, Inoue, A, Wiwatwitaya, D, Ohkuma, M, Kudo, T, Abe, T and Atsuko Sugimoto, A (2005) Carbon mineralization by termites in tropical forests, with emphasis on fungus combs. Ecological Research 20, 453460. CrossRefGoogle Scholar
Zemunik, G, Winter, K and Turner, BL (2020) Toxic effects of soil manganese on tropical trees. Plant & Soil 453, 343354. CrossRefGoogle Scholar
Figure 0

Figure 1. Location of Mae Ping Forest Ecology plot: Dashed line – Mae Ping National Park boundary Shaded – Mae Ping river, Bhumibol Dam, & other lakes; Solid square – Mae Ping Forest Ecology Plot.

Figure 1

Table 1. Land and forest types on Mae Ping 16 ha ecological research plot

Figure 2

Figure 2. Land types, termite mounds, and sampling in the Mae Ping plot Dashed grey line – Termite study area C – Colluvial plain of deep, dystric red loam with dark calcareous clay on termite mounds L – Shallow eutric loam on limestone fins M – Shallow dystric loam on metargillite hillocks Large circle – Mound > 1 m diameter: Filled are sampled; open are unsampled Unsampled smaller mounds: Small circle 0.5 – 1 m diameter; Star < 0.5 m diameter.

Figure 3

Table 2. Termites in mounds on Mae Ping 8 ha study site

Figure 4

Table 3. Forest structure at Mae Ping

Figure 5

Figure 3. Macrotermes annandalei mound in Deciduous Dipterocarp Forest, Mae Ping.

Figure 6

Table 4a. Main species in quadrats in non-mound forest, Mae Ping

Figure 7

Table 4b. Main species in quadrats on termite mounds, Mae Ping

Figure 8

Table 5. Floristic diversity and similarity, Mae Ping

Figure 9

Figure 4. Floristic cluster analyses of quadrats at Mae Ping (a) Trees (b) Saplings (c) Seedlings.

Figure 10

Figure 5. Canonical Correspondence Analyses (CCA) with soil correlations at Mae Ping (a) Trees (b) Saplings (c) Seedlings Open rectangle – Quadrat in non-mound forest; Filled triangle – Quadrat on termite mound; Lines – Pearson correlations of topsoil variable with CCA axis 1.

Figure 11

Table 6. Comparison of non-mound and mound topsoils

Figure 12

Figure 6. Stoichiometric roses of exchangeable cations and pH in Mae Ping topsoils Shaded inner: Non-mound topsoil Clear outer: Mound topsoil All scales, except pH, are linear.

Figure 13

Table 7. Correlations of topsoil variables against CCA axes 1 and 2 for different size classes

Supplementary material: File

Kaewfoo et al. supplementary material

Kaewfoo et al. supplementary material
Download Kaewfoo et al. supplementary material(File)
File 4.7 MB