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Termite-affected soils in Pakistan
0038-0717/82/040359-06$03.00/0 Copyright Q 1982 Pergamon PressLtd
SoilBiol. Biochem.Vol. 14,pp. 359to 364,1982
Printedin Great Britain. All rights reserved
TERMITE-AFFECTED
SOILS IN PAKISTAN
KHALID HAMID SHEIKH and SAFDAR ALI KAYANI*
Department of Botany, Punjab University, New Campus, Lahore, Pakistan (Accepted 20 February 1982) Summary-Soils from mounds and subterranean nests of 13 termite species of Pakistan were analysed and compared with corresponding adjacent termite-free soils. Mound soils had more clay, water-stable aggregates, organic matter, Ca and Mg and higher porosity, water holding and cation exchange capacities than termite-free soils. The magnitude of the changes brought about to soils by the mound builders Odontotermeslokanandi and 0. obesus were almost identical. Subterranean nest soils generally had a greater clay content, higher water holding and cation exchange capacities and greater amounts of Ca and Mg than adjacent termite-free soils. The magnitude of changes between the characteristics of termite-affected and termite-free soils among subterranean nest-building termites varied from species to species. Anacanthotermesmacrocephalusand A. vegans effected maximal changes in soil characteristics while Microcerotermesheimi, M. sukesarensis and Odontotermesgurdaspurensis-brought about minimal changes.
INTRODUCTION constitute an important component of soil fauna in tropical and sub-tropical regions. Pakistan, lying between 24 and 37”N latitude, is mainly subtropical and has a rich termite fauna (Akhtar, 1974). The relative fertility of termite mounds and their adjacent termite-free soils has attracted the attention of pedologists and biologists(Boyer, 1956; Harris, 1956; Hesse, 1955; Maldague, 1959; Stoops, 1964; Wild, 1952; Lee and Wood, 1971a) but subterranean nest soils have received less attention. Lee and Wood (1971a) have analysed the soils from subterranean nests of 5 species of termites and compared them with their adjacent termite-free soils and have found greater amounts of Ca and Mg in the former. We have examined soils from mounds and subterranean nests of 13 different species of termites of Pakistan (Table l), as well as termite-free soils adjacent to these structures, so as to discern changes which termites bring about to soil characteristics.
Termites
METHODS Soil sampling
Samples were collected in various parts of Pakistan (Table 1) from 1973 to 1977. At each site the termiteaffected and corresponding termite-free soils were sampled simultaneously. Only one nest of each species was investigated. Three composite soil samples were obtained from each nest. Termite-affected soils can be divided into two groups based on nest location and structure sampled: mound soil and subterranean nest soil. Soil in a mound was sampled from the central canal which is built entirely of soil cemented with termite saliva. Adjacent termite-free soil was sampled from the depth to which each mound extended, the part from which soil is taken for mound building (Hesse, 1955).
Soils from subterranean nests were sampled from depths which appeared to contain a maximum number of termites (as revealed by a systematic exposure of nest). Adjacent termite-free soils were sampled from corresponding depths. Sampling depths of termite-affected soil in subterranean nests varied from species to species. The subterranean nests of Amitermes belli (Desneux), Coptotermes heimi (Wasmann) and Heterotermes indicoia (Wasmann) were present in decaying stumps of Acacia modesta Wall., Morus alba L. and Pinus roxburghii Sargent, respectively. The stumps were systematically exposed. The termite-affected soils were sampled from the depths at which each stump had a maximum number of termites. Pieces of wood, if present, were removed by hand sorting. Species of Anacanthotermes (A. macrocephalus (Desneux) and A. uagans (Hagen)) build subterranean nests with a very small conical mound on the ground surface. The soils of mounds, galleries and storage chambers were collected. Subterranean nests of Microtermes (M. mycophagus (Desneux), M. obesi Holmgren and M. unicolor Snyder) and Odontotermes gurdaspurensis Holmgren & Holmgren were present, respectively, near the trees of Dalbergia sissoo Roxb. and Acacia arabica (Lam.) Wild. Galleries linked with the main nests, however, were present on the tree trunks. The nests were opened and soils around the fungus combs and those of galleries were sampled as termite-affected soils. Subterranean nests of Microcerotermes heimi Wasmann and M. sakesarensis Ahmad were present in Saccharum munja Roxb. roots. These shallow nests contained a black carton from around which termiteaffected soil was sampled.
* Present address: Department of Botany, Baluchistan University, Quetta, Pakistan. 359
Soil analyses
Soil texture was determined by the hydrometer method of Bouyoucos (1951); water-stable aggregates by “wet sieving” (Richards, 1954); water holding capacity by the “Keen-Raczkowski box” method (Piper,
360
KHALIDHAMIDSHEIKHand SAFDARALI KAYANI Table 1. Sites from which termite-affected
Termite species Odontotermes lokanandi
Sampling site Kotali
and adjacent termite-free soils were sampled in Pakistan
Latitude
Longitude
Vegetation type
33” 03’ N
75” 02’ E
Pinus roxburghii forest
Mango orchard
0. obesus
Lahore
31”30’N
73” 16’ E
Amitermes belli
Jhelum
32” 50’ N
73” 45’ E
Acacia modesta forest
Anacanthotermes ~crocephalus
Bhakhar
31”31’N
?f”04’E
~u~bergia sissoo
A. uagans
Quetta
30” 15’ N
68” 25’ E
irrigated plantation Mixed forest of Pistachia khinjuk & OIea ferruginea
Coptotermes
Chicha-Watni
30” 31’N
D. sissoo
Heterotermes indicala
Murree
33” 5.5’ N
73” 25’ E
irrigated plantation Pinus roxburghii forest
Microtermes mycophagus
Bahawalpur
29” 20’ N
71”40’E
D. sissoo
M. obesi
Bahawatpur
29” 20’ N
71”4O’E
D. sissoo
M. unicolor
Jhang
31”26’N
73” 07’ E
D. sissoo
Microcerotermes heimi
Kot Mithon
28” 25’ N
70” 35’ E
D. sissoo
M. sakesarensis
Lahore
31”30’N
73” 16’E
Road-side plantation of Populus sp.
0. gurdaspurensis
Chicha-Watni
30” 31’N
72” 43’ E
D. sissoo
heimi
irrigated plantation irrigated plantation irrigated plantation irrigated plantation
irrigated plantation
1942); pH with a glass electrode pH meter; organic matter content by the Walkley and Black method (Jackson, 1958); soil cation exchange capacity by leaching with sodium acetate (Richards, 1954); soluble Ca and Mg by titration with ethylenediaminetetr~cetate (EDTA) and soluble Na and K by flame photometry. The results obtained for each of the characteristics of termite-affected and their adjacent soils were subjected to r-test.
RESULTS Soils affected by termites showed markedly different characteristics from their adjacent soils (Tables 2, 3 and 4). bound-building
termites
Mound soils had significantly greater amounts of aggregates and higher clay, more water-stable amounts of soluble Ca and Mg than termite-free soils. Mound soils of Odontotermes lokanandi Chatterjee & Thakur had significantly greater cation exchange capacities while those of 0. obesus (Rambur) had significantly higher electrical conductivities than their corresponding termite-free soils (Tables 2 and 3). Mound soits did not differ significantly from termite-free soils in their water holding capacities, organic matter content, pH and amounts of soluble K and Na. Mean values of water holding capacity and organic matter content, however, were higher in termite-affected soil than in termite-free soil (Tables 2 and 3).
There was no significant difference between mounds built by 0. lokanandi and 0. obesus for various soil characteristics of termite-affected and termite-free soils (Table 4). S~terranean
nest-building
termites
Subterranean nest soils did not differ significantly from termite-free soils with regard to pH, electrical conductivity and soluble Na (Tables 2 and 3). Soils from the nests of Anacanthotermes (A. macrocephalus and A. uagans) had significantly greater cation exchange capacities and organic matter contents than adjacent termite-free soils. Soil of an A. macrocephalus nest also had greater amounts of clay and soluble K than its termite-free counterpart, while soil of an A. uagans nest had greater water holding capacity than its adjoining termite-free soil (Tables 2 and 3). Soils from nests of Amitermes be& and ~optotermes heimi had significantly greater amounts of clay than adjacent termite-free soils. Soil affected by C. heimi also had a significantly higher amount of Ca than the adjacent termite-free soil (Tables 2 and 3). Soils from nests of Microtermes mycophagus, M. obesi and M. unicolor had significantly greater amounts of organic matter, while soil affected by M. unicolor had significantly higher amounts of clay and soluble Ca than the adjacent soil. Soils from nests of M. mycophagus and M. obesi had significantly higher water holding capacity and amount of soluble Mg, respectively, than their corresponding termite-free soils (Tables 2 and 3).
45-60
45-60
Coptotermes heimi
Heterotermes indicola Microtermes mycophagus
30-40
Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent
soil
soil
soil
soii
soil
soil
soil
soil
soil
soil
soil
Mound Adjacent soil Mound Adjacent soil
Site
65.1 tr 1.38 69.3 & 0.0 52.3 k 0.1 56.3 & 1.2
Sand %
63.5 + 0.0 66.3 + 0.0 72.3 rf: 1.3 16.1 + 2.9 83.0 + 5.0 82.6 & 5.2 63.5 zt 1.3 66.4 St 2.9 62.6 3_ 3.2 65.5 k 1.3 65.6 + 2.7 68.4 +_0.0 73.1 & 1.9 15.9 + 3.0 86.1 + 1.3 88.2 +_0.0 78.9 + 2.5 79.3 + 1.5 70.5 + 0.0 71.2 + 1.0 72.3 + 1.3 71.3 + 0.0
* Termite-affected and adjacent soils significantly different at P = 0.05.
0. gurdaspurens~s
O-15
O-15
45-55
M. unicolor
Microcerotermes heimi M. sakesarensis
25-35
M. obesi
30-45
O-15
O-15
30-45
100-115
m-75
Anacanthotermes macrocephalus A. vagans
Subterranean nest-building Amitermes beili
Odontotermes fokanandi 0. obesus
Mound-building
Termite species
Samphng depth cm
+ f f f
0.8 0.0 1.2 0.3
25.6 f 0.0 25.0 + 0.0 19.1 + 0.8 18.8 + 0.6 14.0 f 3.0 14.6 f 3.1 28.4 a. 0.0 28.0 rt: 1.3 21.6 k 1.0 26.5 It 2.0 24.2 f 1.3 23.8 i 0.0 18.3 + 1.3 17.8 * 1.2 7.6 f 2.5 9.1 f 0.0 16.2 f 1.3 15.6 k 1.0 20.9 f 0.0 20.1 + 0.8 22.1 f 0.0 22.9 & 0.0
26.2 25.3 37.9 38.1
Silt %
+_O.O* f 0.0 f l.O* f 1.1
10.9 + o.o* 8.7 +_0.0 8.6 + o.o* 4.5 f 0.3 3.0 + 0.6 2.8 f 0.5 8.1 k 0.5* 5.6 + 0.0 9.8 k 1.3 7.8 f 0.6 10.2 + 0.8 7.8 + 0.0 8.6 f 0.8 6.3 k 0.6 4.4 _+o.o* 2.7 f 0.0 4.8 f 0.8 4.1 f 0.9 8.6 f 0.0 8.8 * 0.2 5.6 f 0.2 5.8 + 0.00
8.7 5.4 9.8 5.6
Clay %
15.8 + 1.6 18.2 t_ 1.6 13.9 + 4.1 13.1 + 1.0 0.6 + 0.1 0.0 + 0.0 6.5 + 0.7 4.8 + 0.5 12.0 + 3.4 15.2 & 2.1 6.7 + 2.1 5.4 + 1.6 3.1 + 0.3 5.0 _t 0.7 0.2 _+0.0 0.3 + 0.0 1.2 + 0.0 1.3 k 0.2 1.6 + 0.1 1.6 + 0.0 3.5 &-0.0 2.8 + 0.2
2.1 +_0.1’ 1.1 k 0.3 3.6 + O.l* 2.1 + 0.1
Water-stable aggregates %
Table 2. Physical characteristics of termite-affected and adjacent termite-free soils. Means and standard errors of three replicates
f + rt f
1.1 1.8 2.8 3.0 38.1 + 1.3 34.6 k 1.5 29.9 + 1.2 22.6 f 1.6 39.1 f 1.3* 32.1 f 0.6 48.0 f 0.8 46.5 + 1.3 54.8 + 1.2 53.0 f 1.8 49.6 f 1.5 41.6 t 1.6 45.1 + 0.0 43.5 + 0.6 41.7 f 0.8 39.8 f 1.8 35.4 i 1.8 33.2 f 1.0 41.1 j, 1.4 39.8 & 0.4 48.5 rt 1.3 46.8 f 1.2
47.0 43.3 51.9 43.2
Water holding capacity %
362
KHALID HAMID SHEIKH and SAFDAR ALI KAYANI
363
Termite-affected soils Table 4. Proportional change* in soil characteristics
Termite species
Clay
Water Cation holding exchange Organic capacity capacity matter
Ca
Mg
Mound-building 0~5~rorermes ~ok~~~ndi 0. obesus Subterranean nest-building Amitermes belli Anacanthotermes macrocephafus A. oagans Coptotermes heimi Heterotermes indicola Microtermes mycophagus M. obesi M. unicolor Microcerotermes heimi M. sakesarensis 0. gurdaspurensis * Proportional
change =
0.61 0.75
0.09 0.20
0.25 0.91 0.07 0.45 0.26 0.31 0.37 0.63 0.02 -0.02 -0.03
0.10 0.32 0.22 0.03 0.03 0.19 0.04 0.05 0.07 0.03 0.04
(Value for termite-aff~ted
0.45 0.17
0.33 0.15
0.55 0.88
0.63 0.64
0.07
0.33
0.13
0.17
0.30 1.24 0.06 0.06 0.30 0.08
1.80 0.47 0.30 0.14 0.24 0.19
2.32 0.38 0.12
2.60 1.17 0.03 0.01
0.10
0.09
0.09
0.33
0.11
0.29 0.11
0.55 0.07
0.07 0.10
0.00 0.07
0.00 0.12
0.00 0.14
0.05 0.03 0.04
0.11
soil) - (Value for termite-free soil)
(Value for termite-free soil)
Soils from nests of Heterotermes indicola, Microcerotermes heimi, M. sakesarensis and Udontotermes gurdaspurensis did not differ markedly from adjacent termite-free soils (Tables 2 and 3). Soil affected by 0. more soluble Mg as compared to the adjacent soil (Table 3). From the point of view of the magnitude of differences between termite-affected soils and their adjacent termite-free soils, the subterranean nest-building termites studied in this work fall into three groups. The first is comprised of Anacanthotermes ~crocephaius and A. uagans whose nest soils differed maximally from adjacent termite-free soils in many of the soil characteristics studied here. The second group includes Amitermes be/Ii, Coptotermes heimi, Heterogurdaspurensis, however, had significantly
termes indicola, Microtermes mycophagus, hf. obesi
and M. unicolor which affected their soil moderately while Microcerotermes heimi, M. sakesarensis and Odontotermes gurdaspurensis constitute the third group. Soils of these termites differed minimally from their adjacent termite-free soils (Table 4). DISCUSSION
A greater amount of clay in mound soils than their adjoining termite-free soils has been reported by Harris (1956), Kemp (1955), Maldague (1959), Nye (1955) and Stoops (1964). Lee and Wood (197ib) have reported that most termite species preferentially select finer particle-size fractions. Their mounds and subterranean nests, therefore, contain more clay and less sand than the adjacent termite-free soils. An increase in organic matter in mound and subterranean nest soils (Tables 3 and 4) is related to the vegetative diet of termites (Bouillon, 1970) and the use of organic materials (salivary or faecal) during nest building. The greater water holding and cation exchange capacities of termite-aff~ted vs adjacent soils are primarily due to their higher contents of clay and organic matter (Tables 2 and 3). The colloidal nature of organic matter is mainly responsible for the higher
water holding capacity of soil; organic matter holds as much as nine times its own weight of water (Daubenmire, 1974). A direct relationship between cation exchange capacity and quantity of organic matter (Kamprath and Welch, 1962; Williams, 1932) and clay (Williams, 1932) has been reported. A greater amount of Ca in mound soils than corresponding termite-free soils has been reported in almost all the studies on this subject (Lee and Wood, 1971b). Higher concentration of Mg in mound soils than in adjacent termite-free soils has been reported by some workers (Boyer, 1956; Pathak and Lehri, 1959; Wild, 1952). Lee and Wood (1971a) have also reported greater amounts of Ca and Mg in subterranean nest soil than in adjacent termite-free soil. An increase in the amounts of Ca and Mg in termiteaffected soils is presumably due to the presence of these elements in plants which are the principal diet of termites. Ca, Mg and K are the principal cations among the mineral constituents of plants and animals (Waksman and Starkey, 1949). Similarly an increase in the amount of K in the soil from nest of Anucanthotermes ~crocepha~us (Table 3) may presumably be due to the higher amount of this element in the plants which serve as its food. There were no marked differences between 0. lokanandi and 0. obesus in the magnitude of differences in characteristics of termite-affected and corresponding termite-free soils (Table 4). The various species of Odontotermes have been reported to have similar mound building behaviour (Hesse, 1955). Moreover, the body sizes of 0. lokanandi and 0. obesus are not very different. It seems that from among the subterranean nest building species studied here, the greater change effected by Anacanthotermes ~crocephalus and A. vagans may be attributed to their relatively much larger body size. Moreover, species of Anacanthotermes are harvester termites. This habit may explain the presence of greater amounts of organic matter in their habitat, which effects changes in other soil
KHALID HAMID SHEIKH and SAFDAR ALI KAYANI
364
characteristics. Relatively small-sized termite species like Amitermes belli, C. heimi, H. indicola, Microtermes mycophagus, M. obesi and M. unicolor effected lesser changes in soil characteristics, while 0. gurdaspurensis, Microcerotermes heimi and M. sakesarensis produced least changes (Table 4). These differences may also be largely due to variations in population density of termites in the sampled nests and differences in the ages of these nests. In the present study, however, no quantitative data have been collected in this regard. Acknowledgements-We thank Professor Dr Muzaffer Ahmad and Dr M. S. Akhtar of Zoology Department, Punjab University, Lahore for their help in the identification of termites. REFERENCES AKHTAR M. S. (1974) Zoogeography of the termites of Pakistan. Pakistan Journal of Zoology 6, 85-104.
BoUILLONA. (1970) Termites of the Ethionian reeion. In Biology of ‘Termites Vol. II, (K. Krishna and F. M. Weesner, Eds), pp. 153-280. Academic Press, New York. Bou~oucos G. J. (1951) A recalibration of the hydrometer method for making mechanical analyses of soils. Aaronomy Journal 43,4%438. BOYER P. (1956) etude oedofoeiaue
de la renartition
et du
dosage hes bases totales da& ies materiaux de la termitiere de Ballicositermes natalensis (Hav.). Comptes rendus hebdomadaire des Seances de I’ Academic des Sciences 242.801-803. DAUBENMIRE R. F. (1974) Plants and Environment, 3rd edn. Wiley, New York. HARRIS W. V. (1956) Termite mound building. Insectes sociaux 3, 261-265. HESSEP. R. (1955) A physical and chemical study of the soils of termite mounds in East Africa. Journal of Ecology 43,449-461.
JACK~CINM. L. (1958) Soil Chemical Analysis. Constable, London. KAMPRATHE. J. and WELCHC. D. (1962) Retention and cation exchange properties of organic matter in Coastal Plains soils. Proceedings Soil Science Society of America 26,263-265.
KEMP P. B. (1955) The termites of north-eastern Tanganyika: their distribution and biology. Bulletin of Entomological Research 46, 113-135. LEE K. E. and WOODT. G. (1971a) Physical and chemical effects on soils of some Australian termites and their pedological significance. Pedobiologia 11, 376409. LEE K. E. and WOOD T. G. (1971b) Termites and Soils. Academic Press, New York. MALDAGUEM. E. (1959) Analyses de sols et materiaux de termititres du Congo Belge. Insectes sociaux 6, 273-288. NYE P. H. (1955) Some soil-forming processes in the humid tropics, IV. The action of soil fauna. Journal of Soil Science 6, 73-83. PATHAKA. N. and LEHRIL. K. (1959) Studies on termite nests. I. Chemical, physical and biological characteristics of a termitarium in relation to its surroundings. Journal of Indian Society of Soil Science 7, 87-90. PIPERC. S. (1942) Soil and P/ant Analysis. Adelaide University, Adelaide. RICHARDSL. A. (Ed.) (1954) Diagnosis and Improvement of Saline and Alkali Soils. USDA Handbook 60, Washington, DC. STOOPSG. (1964) Application of some pedological methods to the analysis of termite mounds. In fitudes sur les Termites Africains. (A. Bouillon, Ed.), pp. 379-398. Leopoldville University, Leopoldville. WAKSMANS. A. and STARKEYR. L. (1949) Soil and Microbes. Wiley, New York. WILD H. (1952) The vegetation of southern Rhodesian termitaria. Rhodesia Agriculture Journal 49, 28@292. WILLIAMSR. (1932) The contribution of clay and organic matter to the base exchange capacity of soils. Journal of Agricultural Science 22, 845-851.
SoilBiol. Biochem.Vol. 14,pp. 359to 364,1982
Printedin Great Britain. All rights reserved
TERMITE-AFFECTED
SOILS IN PAKISTAN
KHALID HAMID SHEIKH and SAFDAR ALI KAYANI*
Department of Botany, Punjab University, New Campus, Lahore, Pakistan (Accepted 20 February 1982) Summary-Soils from mounds and subterranean nests of 13 termite species of Pakistan were analysed and compared with corresponding adjacent termite-free soils. Mound soils had more clay, water-stable aggregates, organic matter, Ca and Mg and higher porosity, water holding and cation exchange capacities than termite-free soils. The magnitude of the changes brought about to soils by the mound builders Odontotermeslokanandi and 0. obesus were almost identical. Subterranean nest soils generally had a greater clay content, higher water holding and cation exchange capacities and greater amounts of Ca and Mg than adjacent termite-free soils. The magnitude of changes between the characteristics of termite-affected and termite-free soils among subterranean nest-building termites varied from species to species. Anacanthotermesmacrocephalusand A. vegans effected maximal changes in soil characteristics while Microcerotermesheimi, M. sukesarensis and Odontotermesgurdaspurensis-brought about minimal changes.
INTRODUCTION constitute an important component of soil fauna in tropical and sub-tropical regions. Pakistan, lying between 24 and 37”N latitude, is mainly subtropical and has a rich termite fauna (Akhtar, 1974). The relative fertility of termite mounds and their adjacent termite-free soils has attracted the attention of pedologists and biologists(Boyer, 1956; Harris, 1956; Hesse, 1955; Maldague, 1959; Stoops, 1964; Wild, 1952; Lee and Wood, 1971a) but subterranean nest soils have received less attention. Lee and Wood (1971a) have analysed the soils from subterranean nests of 5 species of termites and compared them with their adjacent termite-free soils and have found greater amounts of Ca and Mg in the former. We have examined soils from mounds and subterranean nests of 13 different species of termites of Pakistan (Table l), as well as termite-free soils adjacent to these structures, so as to discern changes which termites bring about to soil characteristics.
Termites
METHODS Soil sampling
Samples were collected in various parts of Pakistan (Table 1) from 1973 to 1977. At each site the termiteaffected and corresponding termite-free soils were sampled simultaneously. Only one nest of each species was investigated. Three composite soil samples were obtained from each nest. Termite-affected soils can be divided into two groups based on nest location and structure sampled: mound soil and subterranean nest soil. Soil in a mound was sampled from the central canal which is built entirely of soil cemented with termite saliva. Adjacent termite-free soil was sampled from the depth to which each mound extended, the part from which soil is taken for mound building (Hesse, 1955).
Soils from subterranean nests were sampled from depths which appeared to contain a maximum number of termites (as revealed by a systematic exposure of nest). Adjacent termite-free soils were sampled from corresponding depths. Sampling depths of termite-affected soil in subterranean nests varied from species to species. The subterranean nests of Amitermes belli (Desneux), Coptotermes heimi (Wasmann) and Heterotermes indicoia (Wasmann) were present in decaying stumps of Acacia modesta Wall., Morus alba L. and Pinus roxburghii Sargent, respectively. The stumps were systematically exposed. The termite-affected soils were sampled from the depths at which each stump had a maximum number of termites. Pieces of wood, if present, were removed by hand sorting. Species of Anacanthotermes (A. macrocephalus (Desneux) and A. uagans (Hagen)) build subterranean nests with a very small conical mound on the ground surface. The soils of mounds, galleries and storage chambers were collected. Subterranean nests of Microtermes (M. mycophagus (Desneux), M. obesi Holmgren and M. unicolor Snyder) and Odontotermes gurdaspurensis Holmgren & Holmgren were present, respectively, near the trees of Dalbergia sissoo Roxb. and Acacia arabica (Lam.) Wild. Galleries linked with the main nests, however, were present on the tree trunks. The nests were opened and soils around the fungus combs and those of galleries were sampled as termite-affected soils. Subterranean nests of Microcerotermes heimi Wasmann and M. sakesarensis Ahmad were present in Saccharum munja Roxb. roots. These shallow nests contained a black carton from around which termiteaffected soil was sampled.
* Present address: Department of Botany, Baluchistan University, Quetta, Pakistan. 359
Soil analyses
Soil texture was determined by the hydrometer method of Bouyoucos (1951); water-stable aggregates by “wet sieving” (Richards, 1954); water holding capacity by the “Keen-Raczkowski box” method (Piper,
360
KHALIDHAMIDSHEIKHand SAFDARALI KAYANI Table 1. Sites from which termite-affected
Termite species Odontotermes lokanandi
Sampling site Kotali
and adjacent termite-free soils were sampled in Pakistan
Latitude
Longitude
Vegetation type
33” 03’ N
75” 02’ E
Pinus roxburghii forest
Mango orchard
0. obesus
Lahore
31”30’N
73” 16’ E
Amitermes belli
Jhelum
32” 50’ N
73” 45’ E
Acacia modesta forest
Anacanthotermes ~crocephalus
Bhakhar
31”31’N
?f”04’E
~u~bergia sissoo
A. uagans
Quetta
30” 15’ N
68” 25’ E
irrigated plantation Mixed forest of Pistachia khinjuk & OIea ferruginea
Coptotermes
Chicha-Watni
30” 31’N
D. sissoo
Heterotermes indicala
Murree
33” 5.5’ N
73” 25’ E
irrigated plantation Pinus roxburghii forest
Microtermes mycophagus
Bahawalpur
29” 20’ N
71”40’E
D. sissoo
M. obesi
Bahawatpur
29” 20’ N
71”4O’E
D. sissoo
M. unicolor
Jhang
31”26’N
73” 07’ E
D. sissoo
Microcerotermes heimi
Kot Mithon
28” 25’ N
70” 35’ E
D. sissoo
M. sakesarensis
Lahore
31”30’N
73” 16’E
Road-side plantation of Populus sp.
0. gurdaspurensis
Chicha-Watni
30” 31’N
72” 43’ E
D. sissoo
heimi
irrigated plantation irrigated plantation irrigated plantation irrigated plantation
irrigated plantation
1942); pH with a glass electrode pH meter; organic matter content by the Walkley and Black method (Jackson, 1958); soil cation exchange capacity by leaching with sodium acetate (Richards, 1954); soluble Ca and Mg by titration with ethylenediaminetetr~cetate (EDTA) and soluble Na and K by flame photometry. The results obtained for each of the characteristics of termite-affected and their adjacent soils were subjected to r-test.
RESULTS Soils affected by termites showed markedly different characteristics from their adjacent soils (Tables 2, 3 and 4). bound-building
termites
Mound soils had significantly greater amounts of aggregates and higher clay, more water-stable amounts of soluble Ca and Mg than termite-free soils. Mound soils of Odontotermes lokanandi Chatterjee & Thakur had significantly greater cation exchange capacities while those of 0. obesus (Rambur) had significantly higher electrical conductivities than their corresponding termite-free soils (Tables 2 and 3). Mound soits did not differ significantly from termite-free soils in their water holding capacities, organic matter content, pH and amounts of soluble K and Na. Mean values of water holding capacity and organic matter content, however, were higher in termite-affected soil than in termite-free soil (Tables 2 and 3).
There was no significant difference between mounds built by 0. lokanandi and 0. obesus for various soil characteristics of termite-affected and termite-free soils (Table 4). S~terranean
nest-building
termites
Subterranean nest soils did not differ significantly from termite-free soils with regard to pH, electrical conductivity and soluble Na (Tables 2 and 3). Soils from the nests of Anacanthotermes (A. macrocephalus and A. uagans) had significantly greater cation exchange capacities and organic matter contents than adjacent termite-free soils. Soil of an A. macrocephalus nest also had greater amounts of clay and soluble K than its termite-free counterpart, while soil of an A. uagans nest had greater water holding capacity than its adjoining termite-free soil (Tables 2 and 3). Soils from nests of Amitermes be& and ~optotermes heimi had significantly greater amounts of clay than adjacent termite-free soils. Soil affected by C. heimi also had a significantly higher amount of Ca than the adjacent termite-free soil (Tables 2 and 3). Soils from nests of Microtermes mycophagus, M. obesi and M. unicolor had significantly greater amounts of organic matter, while soil affected by M. unicolor had significantly higher amounts of clay and soluble Ca than the adjacent soil. Soils from nests of M. mycophagus and M. obesi had significantly higher water holding capacity and amount of soluble Mg, respectively, than their corresponding termite-free soils (Tables 2 and 3).
45-60
45-60
Coptotermes heimi
Heterotermes indicola Microtermes mycophagus
30-40
Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent Nest Adjacent
soil
soil
soil
soii
soil
soil
soil
soil
soil
soil
soil
Mound Adjacent soil Mound Adjacent soil
Site
65.1 tr 1.38 69.3 & 0.0 52.3 k 0.1 56.3 & 1.2
Sand %
63.5 + 0.0 66.3 + 0.0 72.3 rf: 1.3 16.1 + 2.9 83.0 + 5.0 82.6 & 5.2 63.5 zt 1.3 66.4 St 2.9 62.6 3_ 3.2 65.5 k 1.3 65.6 + 2.7 68.4 +_0.0 73.1 & 1.9 15.9 + 3.0 86.1 + 1.3 88.2 +_0.0 78.9 + 2.5 79.3 + 1.5 70.5 + 0.0 71.2 + 1.0 72.3 + 1.3 71.3 + 0.0
* Termite-affected and adjacent soils significantly different at P = 0.05.
0. gurdaspurens~s
O-15
O-15
45-55
M. unicolor
Microcerotermes heimi M. sakesarensis
25-35
M. obesi
30-45
O-15
O-15
30-45
100-115
m-75
Anacanthotermes macrocephalus A. vagans
Subterranean nest-building Amitermes beili
Odontotermes fokanandi 0. obesus
Mound-building
Termite species
Samphng depth cm
+ f f f
0.8 0.0 1.2 0.3
25.6 f 0.0 25.0 + 0.0 19.1 + 0.8 18.8 + 0.6 14.0 f 3.0 14.6 f 3.1 28.4 a. 0.0 28.0 rt: 1.3 21.6 k 1.0 26.5 It 2.0 24.2 f 1.3 23.8 i 0.0 18.3 + 1.3 17.8 * 1.2 7.6 f 2.5 9.1 f 0.0 16.2 f 1.3 15.6 k 1.0 20.9 f 0.0 20.1 + 0.8 22.1 f 0.0 22.9 & 0.0
26.2 25.3 37.9 38.1
Silt %
+_O.O* f 0.0 f l.O* f 1.1
10.9 + o.o* 8.7 +_0.0 8.6 + o.o* 4.5 f 0.3 3.0 + 0.6 2.8 f 0.5 8.1 k 0.5* 5.6 + 0.0 9.8 k 1.3 7.8 f 0.6 10.2 + 0.8 7.8 + 0.0 8.6 f 0.8 6.3 k 0.6 4.4 _+o.o* 2.7 f 0.0 4.8 f 0.8 4.1 f 0.9 8.6 f 0.0 8.8 * 0.2 5.6 f 0.2 5.8 + 0.00
8.7 5.4 9.8 5.6
Clay %
15.8 + 1.6 18.2 t_ 1.6 13.9 + 4.1 13.1 + 1.0 0.6 + 0.1 0.0 + 0.0 6.5 + 0.7 4.8 + 0.5 12.0 + 3.4 15.2 & 2.1 6.7 + 2.1 5.4 + 1.6 3.1 + 0.3 5.0 _t 0.7 0.2 _+0.0 0.3 + 0.0 1.2 + 0.0 1.3 k 0.2 1.6 + 0.1 1.6 + 0.0 3.5 &-0.0 2.8 + 0.2
2.1 +_0.1’ 1.1 k 0.3 3.6 + O.l* 2.1 + 0.1
Water-stable aggregates %
Table 2. Physical characteristics of termite-affected and adjacent termite-free soils. Means and standard errors of three replicates
f + rt f
1.1 1.8 2.8 3.0 38.1 + 1.3 34.6 k 1.5 29.9 + 1.2 22.6 f 1.6 39.1 f 1.3* 32.1 f 0.6 48.0 f 0.8 46.5 + 1.3 54.8 + 1.2 53.0 f 1.8 49.6 f 1.5 41.6 t 1.6 45.1 + 0.0 43.5 + 0.6 41.7 f 0.8 39.8 f 1.8 35.4 i 1.8 33.2 f 1.0 41.1 j, 1.4 39.8 & 0.4 48.5 rt 1.3 46.8 f 1.2
47.0 43.3 51.9 43.2
Water holding capacity %
362
KHALID HAMID SHEIKH and SAFDAR ALI KAYANI
363
Termite-affected soils Table 4. Proportional change* in soil characteristics
Termite species
Clay
Water Cation holding exchange Organic capacity capacity matter
Ca
Mg
Mound-building 0~5~rorermes ~ok~~~ndi 0. obesus Subterranean nest-building Amitermes belli Anacanthotermes macrocephafus A. oagans Coptotermes heimi Heterotermes indicola Microtermes mycophagus M. obesi M. unicolor Microcerotermes heimi M. sakesarensis 0. gurdaspurensis * Proportional
change =
0.61 0.75
0.09 0.20
0.25 0.91 0.07 0.45 0.26 0.31 0.37 0.63 0.02 -0.02 -0.03
0.10 0.32 0.22 0.03 0.03 0.19 0.04 0.05 0.07 0.03 0.04
(Value for termite-aff~ted
0.45 0.17
0.33 0.15
0.55 0.88
0.63 0.64
0.07
0.33
0.13
0.17
0.30 1.24 0.06 0.06 0.30 0.08
1.80 0.47 0.30 0.14 0.24 0.19
2.32 0.38 0.12
2.60 1.17 0.03 0.01
0.10
0.09
0.09
0.33
0.11
0.29 0.11
0.55 0.07
0.07 0.10
0.00 0.07
0.00 0.12
0.00 0.14
0.05 0.03 0.04
0.11
soil) - (Value for termite-free soil)
(Value for termite-free soil)
Soils from nests of Heterotermes indicola, Microcerotermes heimi, M. sakesarensis and Udontotermes gurdaspurensis did not differ markedly from adjacent termite-free soils (Tables 2 and 3). Soil affected by 0. more soluble Mg as compared to the adjacent soil (Table 3). From the point of view of the magnitude of differences between termite-affected soils and their adjacent termite-free soils, the subterranean nest-building termites studied in this work fall into three groups. The first is comprised of Anacanthotermes ~crocephaius and A. uagans whose nest soils differed maximally from adjacent termite-free soils in many of the soil characteristics studied here. The second group includes Amitermes be/Ii, Coptotermes heimi, Heterogurdaspurensis, however, had significantly
termes indicola, Microtermes mycophagus, hf. obesi
and M. unicolor which affected their soil moderately while Microcerotermes heimi, M. sakesarensis and Odontotermes gurdaspurensis constitute the third group. Soils of these termites differed minimally from their adjacent termite-free soils (Table 4). DISCUSSION
A greater amount of clay in mound soils than their adjoining termite-free soils has been reported by Harris (1956), Kemp (1955), Maldague (1959), Nye (1955) and Stoops (1964). Lee and Wood (197ib) have reported that most termite species preferentially select finer particle-size fractions. Their mounds and subterranean nests, therefore, contain more clay and less sand than the adjacent termite-free soils. An increase in organic matter in mound and subterranean nest soils (Tables 3 and 4) is related to the vegetative diet of termites (Bouillon, 1970) and the use of organic materials (salivary or faecal) during nest building. The greater water holding and cation exchange capacities of termite-aff~ted vs adjacent soils are primarily due to their higher contents of clay and organic matter (Tables 2 and 3). The colloidal nature of organic matter is mainly responsible for the higher
water holding capacity of soil; organic matter holds as much as nine times its own weight of water (Daubenmire, 1974). A direct relationship between cation exchange capacity and quantity of organic matter (Kamprath and Welch, 1962; Williams, 1932) and clay (Williams, 1932) has been reported. A greater amount of Ca in mound soils than corresponding termite-free soils has been reported in almost all the studies on this subject (Lee and Wood, 1971b). Higher concentration of Mg in mound soils than in adjacent termite-free soils has been reported by some workers (Boyer, 1956; Pathak and Lehri, 1959; Wild, 1952). Lee and Wood (1971a) have also reported greater amounts of Ca and Mg in subterranean nest soil than in adjacent termite-free soil. An increase in the amounts of Ca and Mg in termiteaffected soils is presumably due to the presence of these elements in plants which are the principal diet of termites. Ca, Mg and K are the principal cations among the mineral constituents of plants and animals (Waksman and Starkey, 1949). Similarly an increase in the amount of K in the soil from nest of Anucanthotermes ~crocepha~us (Table 3) may presumably be due to the higher amount of this element in the plants which serve as its food. There were no marked differences between 0. lokanandi and 0. obesus in the magnitude of differences in characteristics of termite-affected and corresponding termite-free soils (Table 4). The various species of Odontotermes have been reported to have similar mound building behaviour (Hesse, 1955). Moreover, the body sizes of 0. lokanandi and 0. obesus are not very different. It seems that from among the subterranean nest building species studied here, the greater change effected by Anacanthotermes ~crocephalus and A. vagans may be attributed to their relatively much larger body size. Moreover, species of Anacanthotermes are harvester termites. This habit may explain the presence of greater amounts of organic matter in their habitat, which effects changes in other soil
KHALID HAMID SHEIKH and SAFDAR ALI KAYANI
364
characteristics. Relatively small-sized termite species like Amitermes belli, C. heimi, H. indicola, Microtermes mycophagus, M. obesi and M. unicolor effected lesser changes in soil characteristics, while 0. gurdaspurensis, Microcerotermes heimi and M. sakesarensis produced least changes (Table 4). These differences may also be largely due to variations in population density of termites in the sampled nests and differences in the ages of these nests. In the present study, however, no quantitative data have been collected in this regard. Acknowledgements-We thank Professor Dr Muzaffer Ahmad and Dr M. S. Akhtar of Zoology Department, Punjab University, Lahore for their help in the identification of termites. REFERENCES AKHTAR M. S. (1974) Zoogeography of the termites of Pakistan. Pakistan Journal of Zoology 6, 85-104.
BoUILLONA. (1970) Termites of the Ethionian reeion. In Biology of ‘Termites Vol. II, (K. Krishna and F. M. Weesner, Eds), pp. 153-280. Academic Press, New York. Bou~oucos G. J. (1951) A recalibration of the hydrometer method for making mechanical analyses of soils. Aaronomy Journal 43,4%438. BOYER P. (1956) etude oedofoeiaue
de la renartition
et du
dosage hes bases totales da& ies materiaux de la termitiere de Ballicositermes natalensis (Hav.). Comptes rendus hebdomadaire des Seances de I’ Academic des Sciences 242.801-803. DAUBENMIRE R. F. (1974) Plants and Environment, 3rd edn. Wiley, New York. HARRIS W. V. (1956) Termite mound building. Insectes sociaux 3, 261-265. HESSEP. R. (1955) A physical and chemical study of the soils of termite mounds in East Africa. Journal of Ecology 43,449-461.
JACK~CINM. L. (1958) Soil Chemical Analysis. Constable, London. KAMPRATHE. J. and WELCHC. D. (1962) Retention and cation exchange properties of organic matter in Coastal Plains soils. Proceedings Soil Science Society of America 26,263-265.
KEMP P. B. (1955) The termites of north-eastern Tanganyika: their distribution and biology. Bulletin of Entomological Research 46, 113-135. LEE K. E. and WOODT. G. (1971a) Physical and chemical effects on soils of some Australian termites and their pedological significance. Pedobiologia 11, 376409. LEE K. E. and WOOD T. G. (1971b) Termites and Soils. Academic Press, New York. MALDAGUEM. E. (1959) Analyses de sols et materiaux de termititres du Congo Belge. Insectes sociaux 6, 273-288. NYE P. H. (1955) Some soil-forming processes in the humid tropics, IV. The action of soil fauna. Journal of Soil Science 6, 73-83. PATHAKA. N. and LEHRIL. K. (1959) Studies on termite nests. I. Chemical, physical and biological characteristics of a termitarium in relation to its surroundings. Journal of Indian Society of Soil Science 7, 87-90. PIPERC. S. (1942) Soil and P/ant Analysis. Adelaide University, Adelaide. RICHARDSL. A. (Ed.) (1954) Diagnosis and Improvement of Saline and Alkali Soils. USDA Handbook 60, Washington, DC. STOOPSG. (1964) Application of some pedological methods to the analysis of termite mounds. In fitudes sur les Termites Africains. (A. Bouillon, Ed.), pp. 379-398. Leopoldville University, Leopoldville. WAKSMANS. A. and STARKEYR. L. (1949) Soil and Microbes. Wiley, New York. WILD H. (1952) The vegetation of southern Rhodesian termitaria. Rhodesia Agriculture Journal 49, 28@292. WILLIAMSR. (1932) The contribution of clay and organic matter to the base exchange capacity of soils. Journal of Agricultural Science 22, 845-851.