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THESIS<br />
THE COMPARATIVE STUDY ON TERMITE DIVERSITY OF MOIST EVERGREEN<br />
FOREST AND DRY EVERGREEN FOREST IN CHANTHABURI PROVINCE<br />
SAJIT CHUTIBHAPAKORN<br />
A Thesis Submitted in Parti<strong>al</strong> Fulfillment of<br />
the Requirements for the Degree of<br />
Master of Science (Agriculture)<br />
Graduate School,<br />
Kas<strong>et</strong>sart University<br />
2002<br />
ISBN 974-357-522-7
ACKNOWLEDGEMENT<br />
The author wishes to express her sincere appreciation and gratitude to Dr.<br />
Charunee Vongk<strong>al</strong>uang, Forest Products Expert, Roy<strong>al</strong> Forest Department, Bangkok,<br />
who is my thesis advisor for her kind guidance and v<strong>al</strong>uable discussion throughout<br />
the course of this study and this thesis. I am grateful to the other members of my<br />
committee, Assistant Professor Dr. Nit Kirtibutr and Associate Professor Dr. Isara<br />
Vongk<strong>al</strong>uang, for their kind encouragement and suggestions of the thesis.<br />
My deep appreciate to Dr. Yupaporn Sornnuwat, the researcher of the Wood<br />
Protection Section, Wood Products Research Division, Roy<strong>al</strong> Forest Department,<br />
Bangkok for her identification <strong>al</strong>l of samples of termites from the study areas and for<br />
her kind advice, and v<strong>al</strong>uable comments and discussions.<br />
I would like to thank the Roy<strong>al</strong> Forest Department for granting permit to<br />
conduct the research in Khao Kitchagoot Nation<strong>al</strong> Park and Khao Soi Dao Wildlife<br />
Sanctuary.<br />
My sincere thank is extended to Dr. Chongrak Wachrinrat and his team from<br />
Department of Silviculture, Faculty of Forestry, Kas<strong>et</strong>sart University for their t<strong>al</strong>ent for<br />
selection of the study sites and plant identification.<br />
Fin<strong>al</strong>ly, I wish to express my gratitude to my parents, my family and other<br />
staffs of the Wood Protection Section, Wood Products Research Division, Roy<strong>al</strong> Forest<br />
Department, Bangkok, for their help and warm encouragement.<br />
Sajit Chutibhapakorn<br />
August 2002
TABLE OF CONTENTS<br />
TABLE OF CONTENTS<br />
LIST OF TABLES<br />
LIST OF FIGURES<br />
INTRODUCTION<br />
Objectives<br />
LITERATURE REVIEW<br />
Termites of Thailand<br />
Classification<br />
Distribution of Termites<br />
Termites in the Tropic<strong>al</strong> Rain Forest<br />
Biology of Termite<br />
The Forest of Thailand<br />
MATERIALS AND METHODS<br />
Materi<strong>al</strong>s<br />
M<strong>et</strong>hods<br />
RESULTS AND DISCUSSION<br />
Specification of Results from Collected Specimens<br />
Diversity of Termite<br />
Dominant Species<br />
Termite Species in Relation to Season<strong>al</strong> Changes<br />
CONCLUSION<br />
LITERATURE CITED<br />
APPENDIX<br />
i<br />
Page<br />
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ii<br />
v<br />
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Table<br />
LIST OF TABLES<br />
1 Termite genera and species recorded in two different forest types :<br />
Moist evergreens forest and dry evergreen forest. MEF = moist<br />
evergreen forest, DEF = dry evergreen forest, F = number of family,<br />
SF = number of subfamily, G = number of genera, SP = number of<br />
species.<br />
2 Termite genera and species recorded in moist evergreen forest and<br />
dry evergreen forest, eastern Thailand classified by their food<br />
habitat. F = number of family, SF = number of subfamily, G =<br />
number of genera, SP = number of species, W = Wood, W&L =<br />
Wood and leaf, S/H = Soil or humus, L = Lichen.<br />
3 Termite genera and species recorded in moist evergreen forest and<br />
dry evergreen forest, eastern Thailand classified by their nest<br />
habitat. F = number of family, SF = number of subfamily, G =<br />
number of genera, SP = number of species, IW = Nest in wood, A =<br />
Arbore<strong>al</strong> nest, E = Epige<strong>al</strong> nest, S = Subterranean nest.<br />
4 Diversity of termites in moist evergreen forest and dry evergreen<br />
forest of Chanthaburi Province, Thailand during cool – dry and hot –<br />
dry season.<br />
ii<br />
Page<br />
37<br />
44<br />
54<br />
64
Table<br />
5 Frequency of termites recorded in each type of forest ecosystem<br />
during cool-dry season and hot-dry season. MEF = moist evergreen<br />
forest, DEF = dry evergreen forest, F = number of family, SF =<br />
number of subfamily, G = number of genera, SP = number of<br />
species.<br />
6 Frequency of dominant species in moist evergreen forest and dry<br />
evergreen forest during cool-dry season and hot-dry season.<br />
7 Number of species of termites recorded in each type of forest<br />
ecosystem during cool-dry season and hot-dry season.<br />
Appendix Table<br />
1 The data recording form used in the field work to d<strong>et</strong>ermine<br />
ecologic<strong>al</strong> niches of termites.<br />
2 Climatic data in moist evergreen forest and dry evergreen forest of<br />
Chanthaburi Province.<br />
3 The volume of water content, percentage of organic matter and soil<br />
litter surface thickness in moist evergreen forest and dry evergreen<br />
forest.<br />
4 The data recording form used in the field work to d<strong>et</strong>ermine<br />
ecologic<strong>al</strong> niches of termites.<br />
iii<br />
Page<br />
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66<br />
70<br />
84<br />
85<br />
85<br />
86
Appendix Table<br />
5 The data recording form used in the field work to d<strong>et</strong>ermine<br />
ecologic<strong>al</strong> niches of termites.<br />
6 Climatic data in moist evergreen forest and dry evergreen forest of<br />
Chanthaburi Province.<br />
iv<br />
Page<br />
87<br />
87
Figure<br />
1 The study site in moist evergreen forest.<br />
2 The study site in dry evergreen forest.<br />
LIST OF FIGURES<br />
3 Model of sampling plots in moist evergreen forest and dry evergreen<br />
forest.<br />
4 Species diversity of termite groups in the study sites (moist<br />
evergreen forest and dry evergreen forest).<br />
5 Species diversity of termite groups in moist evergreen forest (MEF)<br />
and dry evergreen forest (DEF).<br />
6 Species of termites in moist evergreen forest (MEF) and dry<br />
evergreen forest (DEF).<br />
7 Soldier and worker of Angulitermes, newly recorded of termite list of<br />
Thailand. A: Whole body of soldier from above; B: Whole body of<br />
worker from above; C: Whole body of soldier from side.<br />
8 Termite species in differing food habitat in moist evergreen forest<br />
(MEF) and dry evergreen forest (DEF).<br />
9 Wood feeding termites. A: Cryptotermes; B: Microcerotermes<br />
10 Wood and leaf feeding termites. A: Hypotermes; B: Macrotermes<br />
v<br />
Page<br />
28<br />
28<br />
29<br />
30<br />
39<br />
39<br />
42<br />
47<br />
49<br />
50
Figure<br />
11 Soil or humus feeding termites. A: Termes; B: Dicuspiditermes.<br />
12 Lichen feeding termites; Hospit<strong>al</strong>itermes.<br />
13 Termite species in differing nest types in moist evergreen forest<br />
(MEF) and dry evergreen forest (DEF).<br />
14 Wood-dwelling termites. A: Cryptotermes; B: Glyptotermes.<br />
15 Subterranean termites. A: Coptotermes; B: Ancistrotermes.<br />
16 Arbore<strong>al</strong> nesting termites. A: Microcerotermes; B: Nasutitermes.<br />
17 Epige<strong>al</strong> nesting termites. A: Mound of Globitermes; B: Carton nest of<br />
Microcerotermes; C: Mound of Macrotermes; D: Nest of Termes.<br />
18 Frequency of dominant species in moist evergreen forest and dry<br />
evergreen forest during cool-dry season and hot-dry season.<br />
19 Dominant species in moist evergreen forest and dry evergreen<br />
forest. A: Soldier of Microcerotermes crassus (dominant species in<br />
2 forest types); B: Soldier of Globitermes sulphureus (secondary<br />
dominant in moist evergreen forest); C: Soldier of Ancistrotermes<br />
pakestanicus (secondary dominant in moist evergreen forest); D:<br />
Soldier of Schedorhinotermes medioobscurus.<br />
vi<br />
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51<br />
52<br />
58<br />
60<br />
61<br />
62<br />
63<br />
67<br />
69
Figure<br />
20 Number of species of termites recorded in each type of forest<br />
ecosystem during cool-dry season and hot-dry season.<br />
vii<br />
Page<br />
71
THE COMPARATIVE STUDY ON TERMITE DIVERSITY OF MOIST EVERGREEN<br />
FOREST AND DRY EVERGREEN FOREST IN CHANTHABURI PROVINCE.<br />
INTRODUCTION<br />
Thailand is situated within a tropic<strong>al</strong> climatic zone that is governed by monsoon.<br />
The veg<strong>et</strong>ation of Thailand, being under the influence of a monsoon<strong>al</strong> climate is humid<br />
tropic<strong>al</strong> and vast areas are covered with luxuriant forests. Owing to the composite<br />
nature of the physiography, the range of both latitude and longitude, and the variation in<br />
temperature and rainf<strong>al</strong>l the forest of Thailand vary considerably over the country.<br />
Tropic<strong>al</strong> evergreen forests in Thailand are occurred <strong>al</strong>ong the w<strong>et</strong> belt of the<br />
country, usu<strong>al</strong>ly where high rainf<strong>al</strong>l (above 1,500 mm per annum) prevails, and is<br />
strongly effected by the monsoon. It is scattered <strong>al</strong>l over the country either on the<br />
peneplains or the mountains. The tropic<strong>al</strong> evergreen forest is divisible into 3 sub-types<br />
as follow: Tropic<strong>al</strong> rain forest or moist evergreen forest, Dry or semi-evergreen forest and<br />
Hill or lower montane forest (Ne<strong>al</strong>, 1967). Chanthaburi Province has distinguished forest<br />
types that can be counted as representatives of tropic<strong>al</strong> evergreen forest in the studied.<br />
Termites are soil anim<strong>al</strong>s, which are super-abundant in the tropic<strong>al</strong> and<br />
subtropic<strong>al</strong> regions. They can <strong>al</strong>so be found widely spread in some areas of temperate<br />
and occasion<strong>al</strong>ly occur in semi-arid environment (Lee and Wood, 1971; Wood and<br />
Sands, 1978; Swift <strong>et</strong> <strong>al</strong>.,1979; Brian, 1983; Josens, 1985; Wilson, 1990). This<br />
extraordinary abundance of termites is the result of their highly developed soci<strong>al</strong><br />
organization (Myles, 1988; Noirot, 1990; Wilson, 1990; N<strong>al</strong>epa, 1994) and symbiosis with<br />
microorganism (Martin, 1987; Wood and Thomas, 1989; Breznak and Brune, 1994).<br />
Termites play an important role of super-decomposer (Matsumoto and Abe, 1979;<br />
Collins, 1981; 1983; Martius, 1994) and carbon-nitrogen b<strong>al</strong>ancer in the tropic<strong>al</strong><br />
terrestri<strong>al</strong> ecosystems of which they are a biotic constituent (Higashi <strong>et</strong> <strong>al</strong>., 1992), thus<br />
forming the basis for a large food web (Deligne <strong>et</strong> <strong>al</strong>., 1981).
Termites are <strong>al</strong>so named as ecosystem engineers (Jones <strong>et</strong> <strong>al</strong>., 1994; Lawton,<br />
1994) that modify the soil structure by constructing mounds and subterranean nests<br />
(Lee and Wood, 1971; Wood, 1988) providing many species of anim<strong>al</strong>s and plants with<br />
diverse habitats (Glover <strong>et</strong> <strong>al</strong>., 1964).<br />
Termites are widely distributed in tropic<strong>al</strong> and subtropic<strong>al</strong> regions. The number<br />
of species and their biomass are especi<strong>al</strong>ly large in the tropic<strong>al</strong> zone. In a wide range<br />
of tropic<strong>al</strong> climates, from savanna to rain forest, termites are believed to play an<br />
important role on the turnover of organic matter and the maintenance and improvement<br />
of soil fertility (Krishna and Weesner, 1969; 1970; Lee and Wood, 1971).<br />
Some important studies were carried out on the abundance, distribution, feeding<br />
habits and role of termites in the savannah ecosystem of West Africa (Wood <strong>et</strong> <strong>al</strong>., 1977)<br />
and in lowland rain forest of west M<strong>al</strong>aysia (Abe and Matsumoto, 1979). In Thailand<br />
Ahmad (1965) described 74 species of termites over the whole area of the country.<br />
Morimoto (1973) presented paper de<strong>al</strong>s with 90 species, of which four are new to<br />
science and 13 are new to Thailand. Intanai (1987) recorded 25 species of termites in<br />
robber plantation of Chanthaburi and Trat provinces which two species, Hypotermes<br />
obcuriceps and Nasutitermes profuscipennis were the new record of Thailand. Davies<br />
(1997) surveyed termite species richness in 2 types of dry deciduous dipterocarp forest<br />
in Doi-Suthep-Pui Nation<strong>al</strong> Park. The result recorded 10 and 13 termite species in fireprotected<br />
and non fire-protected site. However no information has been available on the<br />
termites diversity in moist evergreen forest and dry evergreen forest and the ecosystem<br />
which would govern the existing termites diversity in this part of the world especi<strong>al</strong>ly in<br />
Chanthaburi Province which represents eastern Thailand for the mentioned forest types.<br />
2
Objectives<br />
1. To survey and study the diversity of termites in moist evergreen forest and<br />
dry evergreen forest.<br />
2. To distinguish and characterize specific characters of forest types which<br />
influence species diversity of termite in the study areas.<br />
3. To prepare scientific note on ecologic<strong>al</strong> nitches of termite species collected<br />
from the survey.<br />
3
LITERATURE REVIEW<br />
Termites of Thailand<br />
Termites of Thailand had been very poorly investigated until the excellent<br />
monograph made by Ahmad (1965). Holmgren (1913) first recorded five species of<br />
termites of Thailand and Snyder (1949) listed six species in his cat<strong>al</strong>ogue of the termites<br />
of the world. Ahmad’s monograph contains 74 species and 19 species were<br />
enumerated by Harris (1968). Morimoto (1973) presented paper de<strong>al</strong>s with 90 species,<br />
of which four are new to science and 13 are new to fauna of Thailand. Watanabe <strong>et</strong> <strong>al</strong>.<br />
(1984) listed 16 species of termites at Nam Prom, northeast of Thailand and their types<br />
of mound and nest are described. Intanai (1987) found 25 species of termites belonging<br />
to 13 genera in 6 subfamilies and 3 families which the following termites Hypotermes<br />
obcuriceps and Nasutitermes profuscipennis were the first record in Thailand.<br />
Altog<strong>et</strong>her, 92 species of termites have been liter<strong>al</strong>ly recorded from Thailand up to the<br />
present.<br />
Classification<br />
In the anim<strong>al</strong> kingdom termites are placed in a separate order of the class<br />
Insecta, a order Isoptera which mean “equ<strong>al</strong>-winged” (Isos in Greek meaning the same<br />
and ptero meaning wings) on account that the winged adults having two pairs of wings<br />
of similar shape and size (Hickin,1971; Eaton and H<strong>al</strong>e, 1993).<br />
Termites, just the same as ants, have castes such as workers, soldiers and<br />
winged reproductives. Due to their poor abdomin<strong>al</strong> sclerization, especi<strong>al</strong>ly in the worker<br />
caste, they appear white and are usu<strong>al</strong>ly often c<strong>al</strong>led “white ants” in most part of the<br />
world (Pearce, 1999). But this common name is a misinterpr<strong>et</strong> since they are only<br />
distantly related to the true ants or Formicidae, though the two groups offer striking<br />
an<strong>al</strong>ogies in structure and habits (Wilson, 1971).<br />
4
Snyder (1948) gave some of the other vernacular names for termites such as<br />
“wood lice”, “hormigas blancas”, “comejen”, “cupim”, “carcoma” and “polilla” in the<br />
Americas; “weisse Ameisen” and “fourmis blanches” in Europe; “anay” in the<br />
Philippines; and “Rysmiere” in South Africa.<br />
All of the termites tog<strong>et</strong>her comprise the Order Isoptera. Sever<strong>al</strong> versions of<br />
higher classification, dividing the order into either five or six families, have been offered<br />
in recent years (Snyder, 1948; 1949; Krishna, 1969; Wilson, 1971). Emerson (1965)<br />
recognized six families including Mastotermitidae, K<strong>al</strong>otermitidae, Hodotermitidae,<br />
Rhinotermitidae, Serritermitidae, and Termitidae. Higashi and Abe (1997); Pearce<br />
(1999); Eggl<strong>et</strong>on (2000) reported termite of the world to be consisted of about 2,200<br />
species divided into two large groups: the lower termites (Mastotermitidae,<br />
K<strong>al</strong>otermitidae, Termopsidae, Hodotermitidae, Rhinotermitidae, and Serritermitidae) and<br />
the higher termites (Termitidae), and seven families which includes:<br />
1. Family Mastotermitidae<br />
This group has one extant member, Mastotermes darwiniensis Froggatt, found<br />
only in non-rain forest parts of northern Austr<strong>al</strong>ia (Watson and Abbey, 1993).<br />
2. Family Hodotermitidae<br />
This group are an entirely Old World group of surface–foraging, grass–feeding<br />
termites, found in habitats ranging from dry savanna to arid grassland (Johnson and<br />
Wood, 1980).<br />
3. Family Termopsidae<br />
All had known species of this family nest and feed in rotting logs.<br />
5
4. Family K<strong>al</strong>otermitidae<br />
This family are referred to as dry-wood termites and includes the damp-wood<br />
and powder-post termites (Borrer and Delong, 1971; Wilson, 1971) because they nest<br />
primarily in wood, without soil connection. They were described to have evolved from<br />
the Mastotermitidae (Krishna, 1970). Richards and Davies (1977) reported that this<br />
family has over 250 species and Neotermes, K<strong>al</strong>otermes and Glytotermes are the<br />
important genera.<br />
5. Family Rhinotermitidae<br />
Eggl<strong>et</strong>on (2000) described that this family are <strong>al</strong>l wood feeders and are<br />
distributed widely across tropic<strong>al</strong>, subtropic<strong>al</strong> and temperate regions <strong>al</strong>though there are<br />
no endemic genera in Africa. They <strong>al</strong>so inhabit a very wide range of habitats from w<strong>et</strong> to<br />
dry. Krishna (1970) divided this family into six subfamilies, <strong>al</strong>most <strong>al</strong>l species are<br />
subterranean termites and Coptotermes and R<strong>et</strong>iculitermes are major pests in Asia,<br />
America and Europe.<br />
6. Family Serritermitidae<br />
Richards and Davies (1977) described that this family includes only Serritermes<br />
serrifer from Brazil, with unusu<strong>al</strong> f<strong>al</strong>cate mandibles in adult and worker. This species has<br />
been variously placed in the K<strong>al</strong>otermitidae, Rhinotermitidae and Termitidae.<br />
7. Family Termitidae<br />
This is the largest family of termites and consists of three-quarters of <strong>al</strong>l known<br />
species (Krishna, 1970; Richards and Davies, 1977). Krishna (1970) grouped this family<br />
into 4 subfamilies; they are Amitermitinae, Termitinae, Macrotermitinae and<br />
6
Nasutitermitinae. Watson and Gay (1997) divided this family into 4 subfamilies; they are<br />
Macrotermitinae, Apicotermitinae, Termitinae and Nasutitermitinae.<br />
7.1 Subfamily Macrotermitinae<br />
Darlington (1994) described that species within this subfamily, the fungusgrowing<br />
termites, feed on a wide range of dead and living plant materi<strong>al</strong> (e.g. grass, leaf<br />
litter, or wood) that is fully processed by their fung<strong>al</strong> symbiont (the basidiomyc<strong>et</strong>e,<br />
Termitomyces) on fungus combs in the nest.<br />
7.2 Subfamily Nasutitermitinae<br />
Higashi and Abe (1997) explained that the Nasutitermitinae, which contain<br />
more than 500 species nest in various sites such as in dead wood, on tree trunks, on the<br />
ground, and in soil, and they consume various dead plant materi<strong>al</strong>s such as dead wood,<br />
dead grasses, soil, dung, and <strong>al</strong>so lichen. Nasutitermitinae commonly built their nests on<br />
very high trees over 5-10 m<strong>et</strong>ers (arbore<strong>al</strong> nest) and feed on semi-decomposed leaf and<br />
twigs in contact with soil or dead branches.<br />
7.3 Subfamily Amitermitinae<br />
Roonw<strong>al</strong> (1970) described that this subfamily is characterized by soldiers<br />
with a subrectangular head gener<strong>al</strong>ly having saber-shaped mandibles with a single<br />
fairly prominent tooth or with the margin more or less serrated.<br />
7.4 Subfamily Termitinae<br />
Eggl<strong>et</strong>on (2000) described that this group has characteristic<strong>al</strong>ly<br />
asymm<strong>et</strong>ric<strong>al</strong> mandibles in the soldiers and gener<strong>al</strong>ly either soil-wood interface feeders<br />
7
or soil feeders. They are found throughout the tropics, but are most diverse in tropic<strong>al</strong><br />
Austr<strong>al</strong>ia (Miller, 1991) and tropic<strong>al</strong> forests in the M<strong>al</strong>ay Archipelago (Eggl<strong>et</strong>on, 1997).<br />
7.5 Subfamily Apicotermitinae<br />
This termites are soldierless soil feeders that form diffuse underground nests<br />
in the soil (Eggl<strong>et</strong>on, 2000).<br />
The lower termites (Mastotermitidae, K<strong>al</strong>otermitidae, Termopsidae,<br />
Hodotermitidae, Rhinotermitidae, and Serritermitidae) are characterized by the presence<br />
of cellulolytic protozoa in their hindgut, and mainly consume wood, while the higher<br />
termites (Termitidae) are characterized by the absence of those protozoa, and variously<br />
consume a range of dead and decaying plant materi<strong>al</strong> including sound wood, standing<br />
and f<strong>al</strong>len plant shoots and leaves, decaying wood, and soil (Wood, 1978).<br />
According to their mode of living, termites were found to dwell in a vari<strong>et</strong>y of<br />
habitats which are mostly shut out from light, i.e. in soil, inside wood, under cow dung, in<br />
arbore<strong>al</strong> nests, <strong>et</strong>c. Roonw<strong>al</strong> (1970) grouped them as follows:<br />
1. Wood-dwellers<br />
The colony of wood-dwellers is confined throughout its life to wood, which is<br />
usu<strong>al</strong>ly above ground. This wood may be living or dead, which the swarming pair enters<br />
to start a new colony.<br />
- Dry-wood termites are able to flourish in dry, sound, seasoned wood.<br />
- Damp-wood termites require wood with a constant high amount of<br />
moisture as is to be found in dead or decaying logs and in tree-stumps in moist, cool<br />
regions.<br />
8
2. Ground-dwellers<br />
The colony of ground dwelling termites <strong>al</strong>ways lives at least partly in the<br />
ground; the colonizing pair enter the earth or wood lying in the earth. Ground dweller<br />
consists of 3 subgroups.<br />
- Subterranean termites live underground, but build covered runways to<br />
reach the wood above ground. Nests are built either in the soil underground or in wood<br />
above ground.<br />
- Mound builders construct conspicuous earthen mounds, which may<br />
reach a height of about 3 m above the ground in the Orient and have characteristic<br />
shapes varying with the species.<br />
- Carton-nest-builders construct cellular nests of carton (a substance<br />
made of a mixture of termite fec<strong>al</strong> matter, woody fragments, and som<strong>et</strong>imes earth) which<br />
may be papery and fragile or woody and hard so as to require an axe to break open.<br />
The carton nests may be arbore<strong>al</strong> or partly or wholly buried in the earth.<br />
Distribution of Termites<br />
Snyder (1948) explained that termites are widely distributed throughout the<br />
word, and living species occur in <strong>al</strong>l of the zoogeographic<strong>al</strong> regions except the Arctic<br />
and Antarctic. At present, the African or Ethiopian region is richest in number of species;<br />
the Orient<strong>al</strong> fauna includes the next greatest number of kinds; then come the<br />
Neotropic<strong>al</strong> or tropic<strong>al</strong> American and Austr<strong>al</strong>ian zoogeographic<strong>al</strong> regions with large<br />
numbers of termite species; and fin<strong>al</strong>ly the North American or Nearctic and P<strong>al</strong>aearctic<br />
termite fauna, with relatively few different kinds. Species are most prev<strong>al</strong>ent in the<br />
regions where they originated; primitive kinds remain, and advanced species migrate.<br />
9
Collins (1997); Pearce (1999) explained that the world distribution of termites is<br />
constrained primarily by temperature and moisture.<br />
Krishna (1970) gave a unique combination of factors affecting the distribution of<br />
termites, such as:<br />
1. The great majority of termites can live only under tropic<strong>al</strong> conditions, and<br />
only a very few species have successfully invaded cold climates.<br />
2. Termites live on a restricted di<strong>et</strong> of cellulose (wood, leaves, and humus) and<br />
have not been able to radiate adaptively into various food niches.<br />
3. Termites are soft bodied insects which much as a consequence live in a<br />
closed environment, the nest, with regulated temperature and humidity, which limited in<br />
their mobility.<br />
4. Being soci<strong>al</strong> insect insects, termites are virtu<strong>al</strong>ly limited in their means of<br />
dispers<strong>al</strong> to the swarming period.<br />
5. Even during swarming, termites are weak fliers and are not likely to disperse<br />
over wide water gaps; mating occurs only after flight, and the likelihood is slim that both<br />
a m<strong>al</strong>e and a fem<strong>al</strong>e would be carried by wind over a long distance and land never<br />
each other.<br />
6. Termites are more vulnerable to predators than many other groups of<br />
anim<strong>al</strong>s.<br />
Termites in the Tropic<strong>al</strong> Rain Forest<br />
Tropic<strong>al</strong> rain forests are the stronghold of the Isoptera. The Mastotermitidae,<br />
Termopsidae and Hodotermitidae do not occur in rain forest. The Serritermitidae<br />
includes only one little known species found in wooded savannas in Brazil (Emerson and<br />
Krishna, 1975).<br />
10
The K<strong>al</strong>otermitidae (dry wood termites) are often difficult to find, but probably<br />
occur in sm<strong>al</strong>l numbers in <strong>al</strong>l rain forests. They may <strong>al</strong>so be found on the forest floor<br />
after tree f<strong>al</strong>ls.<br />
The most important family of lower termites in rain forests is the Rhinotermitidae<br />
(damp wood termites), which probably evolved in the Orient<strong>al</strong> region (Emerson, 1955).<br />
Over 73% of termite species are in the higher termite family Termitidae and in terms of<br />
species richness and population density they predominate in <strong>al</strong>l tropic<strong>al</strong> rain forests<br />
(Collins, 1997).<br />
Termites have an impact on rain-forest ecosystems in three main ways, with a<br />
fourth possibility poorly understood (Collins, 1997). They are:<br />
1. feeding on plant materi<strong>al</strong>, often of a very poor qu<strong>al</strong>ity;<br />
2. reproducing and providing materi<strong>al</strong> for predatory food chains;<br />
3. physic<strong>al</strong>ly translocating and chemic<strong>al</strong>ly <strong>al</strong>tering soils through their building<br />
activities; and<br />
4. fixing atmospheric nitrogen.<br />
All of these factors are related to the abundance and biomass of termites in<br />
forests, measurement of which is fundament<strong>al</strong> to a greater understanding of termite<br />
ecology.<br />
Biology of Termite<br />
All termites are eusoci<strong>al</strong> insect (Wilson, 1971). They are polymorphic and live in<br />
large communities (Hickin, 1971; Richard and Davies, 1977). There are well over 2,500<br />
different species of termites in the world (Pearce, 1999).<br />
11
Termites are som<strong>et</strong>imes c<strong>al</strong>led white ants, but they differ from the ants in sever<strong>al</strong><br />
ways. Termites are very soft-bodied and usu<strong>al</strong>ly light-colored, while ants are hardbodied<br />
and usu<strong>al</strong>ly dark-colored; the front and hind wings of a termite are similar in size<br />
and venation and are held flat over the abdomen at rest, but in ants the hind wings are<br />
sm<strong>al</strong>ler than the front wings and have fewer veins, and the wings at rest are usu<strong>al</strong>ly held<br />
above the body. The abdomen in termites is broadly joined to the thorax, whereas in<br />
ants it is constricted at the base and connected to the thorax by a narrow p<strong>et</strong>iole. The<br />
antennae of a termite are moniliform or filiform, while those of ants are elbowed (Borror<br />
and Delong, 1971).<br />
1. Extern<strong>al</strong> morphology (Weesner, 1969)<br />
The Isoptera have three distinct body regions : head, thorax, and abdomen.<br />
1.1 The head: Important structures located on the head which are used for<br />
identification are:<br />
1.1.1 Dors<strong>al</strong> and later<strong>al</strong> structures<br />
a. The head capsule.<br />
b. The labrum.<br />
c. The clypeus.<br />
d. The antennae.<br />
e. The compound eyes.<br />
f. The ocelli.<br />
g. The fontanelle. A sm<strong>al</strong>l opening may be observed in the the<br />
midline on the top of head, just behind or b<strong>et</strong>ween the compound eyes.<br />
1.1.2 Ventr<strong>al</strong> structures<br />
a. The mandibles.<br />
b. The maxillae.<br />
12
c. The labium.<br />
1.2 The thorax: The thorax is a complex structure with three distinct<br />
segments: the prothorax (anteriorly), the mesothorax, and the m<strong>et</strong>athorax, each bearing<br />
a pair of legs. The mesothorax and m<strong>et</strong>athorax <strong>al</strong>so bear a pair of wings in the <strong>al</strong>ate.<br />
1.3 The abdomen: The abdomen includes ten segments with a s<strong>et</strong> of<br />
sclerotized plates: tergite, above, and sternites, below.<br />
2. The castes of termites<br />
Caste d<strong>et</strong>ermination in the lower termites is based primarily on pheromones ; in<br />
some of the higher termites it involves sex ; but the other factors remain unidentified<br />
(Wilson, 1971).<br />
A termite colony consists of three castes : workers or pseudoworker, soldiers<br />
and reproductives (Krishna, 1969; Wilson, 1971; Pearce, 1999).<br />
queens.<br />
2.1 Reproductives (<strong>al</strong>ates or swarmers)<br />
These are the winged reproductives which can produce new kings and<br />
2.1.1 Primary Reproductives<br />
The members of this caste have two pairs of large membranous<br />
wings which are shed after they have swarmed in the process of founding new colonies.<br />
Their body is well sclerotized and more or less dark brown in colour, compound eyes<br />
are fully developed and there are often paired ocelli. Primary reproductives are adapted<br />
for a short aeri<strong>al</strong> life, after which they pair and form the king and queen of a new colony.<br />
13
2.1.2 Supplementary Reproductives<br />
The members of this caste have no aeri<strong>al</strong> life, the body is usu<strong>al</strong>ly<br />
much less sclerotized and pigmented than in the primary reproductives and the<br />
compound eyes are gener<strong>al</strong>ly reduced. Growth of the wings is inhibited to varying<br />
degree and they usu<strong>al</strong>ly resemble wing-pads, though with some indications of venation.<br />
The brain, front<strong>al</strong> gland and reproductive system are <strong>al</strong>so somewhat reduced in size.<br />
Some authorities distinguish three kinds of supplementary reproductives: adultoid<br />
supplementaries very similar in appearance to the primaries, nymphoid supplementaries<br />
with short, pad-like wings, and ergatoid supplementaries without any wing rudiment.<br />
Supplementary reproductives are not norm<strong>al</strong>ly found in colonies<br />
headed by the origin<strong>al</strong> primary reproductives.<br />
2.2 Worker<br />
True workers are absent from some termites, their place being taken by<br />
nymph<strong>al</strong> stages or pseudergates. Where they occur, however, worker are numeric<strong>al</strong>ly<br />
the most important members of the community. They are usu<strong>al</strong>ly p<strong>al</strong>e in colour, with the<br />
cuticle only slightly sclerotized, and they look more like nymphs than the adults of the<br />
other castes. Extern<strong>al</strong> sexu<strong>al</strong> characters are hardly perceptible though workers may be<br />
gen<strong>et</strong>ic<strong>al</strong>ly m<strong>al</strong>e or fem<strong>al</strong>e. The head is directed downwards; it is relatively wider than in<br />
the reproductive castes but never as large as in soldiers. Compound eyes are<br />
reasonably well-developed in the workers of the Hodotermitidae that forage above<br />
ground in daylight, but are otherwise vestigi<strong>al</strong> or absent. The thorax resembles that of<br />
soldiers rather than reproductives. Specific characters are not clearly expressed and<br />
unassociated worker termites may be difficult or impossible to identify.<br />
They exhibit marked care for the eggs and young and in times of danger<br />
may remove them to safer situations. They <strong>al</strong>so feed and tend the queens, forage for<br />
14
food, often at a distance from the nest and, in the fungus-growing species, attend to the<br />
cultivation of the “garden”.<br />
2.3 Soldiers<br />
There are structur<strong>al</strong>ly the most speci<strong>al</strong>ized members of the community, but<br />
their occurrence in virtu<strong>al</strong>ly <strong>al</strong>l genera and certain features of intercastes suggest that<br />
they are the primitive sterile caste. Four well-defined types of soldiers can be<br />
distinguished:<br />
2.3.1 mandibulate soldiers, the most frequent form, with large, powerful<br />
mandibles that may assume striking or grotesque shapes;<br />
2.3.2 nasute soldiers, in which the mandibles are vestigi<strong>al</strong> but there is a<br />
long median front<strong>al</strong> nasus or rostrum, at the end of which the front<strong>al</strong> gland openes; they<br />
occur only in the Nasutitermitinae;<br />
2.3.3 nasutoid soldiers, found among the Rhinotermitidae, produced into a<br />
snout-like structure bearing a dors<strong>al</strong> channel <strong>al</strong>ong which the secr<strong>et</strong>ion of the front<strong>al</strong><br />
gland can flow;<br />
2.3.4 phragmotic soldiers, found in the K<strong>al</strong>otermitidae, whose head is a<br />
strongly sclerotized, more or less plug-like structure that can be used to block the nest<br />
openings and whose mandibles are reduced.<br />
Like the workers, soldiers usu<strong>al</strong>ly consist of gen<strong>et</strong>ic m<strong>al</strong>es and fem<strong>al</strong>es, but<br />
in some Termitidae caste and sex-d<strong>et</strong>ermination seem to be associated : most<br />
Nasutitermitinae have only m<strong>al</strong>e soldiers while in the Macrotermitinae and Termitinae<br />
they are fem<strong>al</strong>e.<br />
15
Soldiers are speci<strong>al</strong>ized in structure and behavior for the defence of the<br />
colony, mainly against other insects-espici<strong>al</strong>ly ants- and vertebrate predators.<br />
3. The termites community and its biology<br />
3.1 Colony-founding<br />
The typic<strong>al</strong> m<strong>et</strong>hod of founding new colonies is by the emission of swarms<br />
of primary reproductives. M<strong>al</strong>es and fem<strong>al</strong>es leave the nest in about equ<strong>al</strong> numbers<br />
and may emerge in a continuous stream or a series of sm<strong>al</strong>ler batches. Swarming may<br />
be diurn<strong>al</strong> or nocturn<strong>al</strong> according to the species, those which swarm at night being<br />
frequently attracted to light. Swarming is often a season<strong>al</strong> occurrence of limited duration<br />
or may occur at times over a rather long period e.g. many tropic<strong>al</strong> species do so after<br />
the first rains of the rainy season. After leaving the nest the <strong>al</strong>ates fly weakly for a short<br />
distance.<br />
Attacks by birds, lizards and sm<strong>al</strong>l mamm<strong>al</strong>s cause a high mort<strong>al</strong>ity among<br />
swarming <strong>al</strong>ates but the survivors, on compl<strong>et</strong>ing their flight, cast their wings and on<br />
encountering a de<strong>al</strong>ate of the opposite sex show some epigamic behavior before<br />
w<strong>al</strong>king in tandem fashion (the m<strong>al</strong>e following the fem<strong>al</strong>e) to seek a site where, in wood<br />
or the ground, they excavate a sm<strong>al</strong>l nupti<strong>al</strong> chamber in which copulation takes place.<br />
At least in the higher termites, the whole series of acts beginning with the emission of the<br />
swarm and ending with the establishment of the roy<strong>al</strong> pair in the nupti<strong>al</strong> chamber are<br />
closely linked tog<strong>et</strong>her.<br />
3.2 Growth of the colony<br />
Egg-production by the primary queen is low at first 15 - 50 eggs may be laid<br />
in the first season, of which some are eaten by the parents. Later, the fecundity<br />
increases and mature queens of the Termitidae lay sever<strong>al</strong> thousand eggs per day (at<br />
16
least over short periods) though far fewer are produced by queens of the primitive<br />
families. The size attained by a mature colony is much sm<strong>al</strong>ler in the primitive genera.<br />
On the other hand, colonies of the Termitidae may contain over a million individu<strong>al</strong>s, <strong>al</strong>l<br />
derived from a single roy<strong>al</strong> pair. The individu<strong>al</strong>s produced in the early stages of a<br />
colony are <strong>al</strong>ways of the sterile castes; <strong>al</strong>ate reproductives develop later. Member of the<br />
sterile castes probably live for 2-4 years. The longevity of the reproductive forms is not<br />
know accurately; it may be 15-50 years in the higher forms but is shorter in primitive<br />
families.<br />
The Forest of Thailand<br />
The climate of Thailand is humid tropic<strong>al</strong>, being influence by season<strong>al</strong> monsoon<br />
winds and by the loc<strong>al</strong> topography. M<strong>et</strong>eorologic<strong>al</strong>ly, two types of climate are<br />
recognized, that of the tropic<strong>al</strong> rain forest and that of the tropic<strong>al</strong> Savannah. The tropic<strong>al</strong><br />
rain forest climate is characterized by uniformly high temperatures and heavy rainf<strong>al</strong>l,<br />
and by the absence of a distinct dry season. The Tropic<strong>al</strong> Savannah climate has less<br />
rainf<strong>al</strong>l and comprises three seasons, cool dry, hot dry, and rainy. The cool dry season<br />
extends from November to February, the hot dry season extends through March and<br />
April and the rainy season covers the longest period from May to October.<br />
The veg<strong>et</strong>ation of Thailand, being under the influence of a monsoon climate is<br />
humid tropic<strong>al</strong> and vast areas are covers with luxuriant forests. Owing to the composite<br />
nature of the physiography, the range of both latitude and longitude, and the variation in<br />
temperature and rainf<strong>al</strong>l the forest of Thailand vary considerably over the country<br />
(Siripornnoppakun, 1989).<br />
The forest of Thailand can be divided into two primary categories, which is<br />
Evergreen and Deciduous.<br />
1. Evergreen Forest<br />
17
The evergreen forest is composed of a high proportion of species lacking a<br />
leafless period. It can be subdivided into the four following type:<br />
1.1 Tropic<strong>al</strong> Evergreen Forest<br />
1.2 Coniferous Forest<br />
1.3 Swamp Forest<br />
1.4 Beach Forest<br />
1.1 Tropic<strong>al</strong> Evergreen Forest<br />
This type of forest occurs <strong>al</strong>ong the w<strong>et</strong> belt of the country, usu<strong>al</strong>ly where<br />
high rainf<strong>al</strong>l (above 1,500 mm per annum) prevails, and is strongly affected by the<br />
monsoon. It is scattered <strong>al</strong>l over the country either on the peneplains or the mountains.<br />
The tropic<strong>al</strong> evergreen forest is again divisible into three sub-types as follow:<br />
1.1.1 Tropic<strong>al</strong> Rain Forest<br />
1.1.2 Dry or Semi-Evergreen Forest<br />
1.1.3 Hill or Lower Mountain Forest<br />
1.1.1 Tropic<strong>al</strong> Rain Forest<br />
In the Chanthaburi or Southeastern and Peninsular Regions, where<br />
impact of the monsoon is directly, the rainf<strong>al</strong>l is very high (2500 mm up), and for this<br />
reason nearly the whole of this area is covered with this type of forest.<br />
The lower tropic<strong>al</strong> rain forest occupies the peneplains and<br />
hillslopes up to 600 m <strong>al</strong>titude. The forests so occurred are being two storied, the upper<br />
story is composed of gigantic trees mostly of Dipterocarps (Dipterocarpus, Hopea,<br />
Shorea, B<strong>al</strong>lanocarpus, Parashorea, and Anisoptera), and other such as Dyera,<br />
18
Endospermum, Melanorrhoea, Mangifera, Swintonia, <strong>et</strong>c. The lower story is constituted<br />
of tree of medium height and girth of genera Vatica, T<strong>al</strong>auma, Macaranga, M<strong>al</strong>lotus,<br />
Glochidon, Syzygium, <strong>et</strong>c. P<strong>al</strong>m and vines are abundant, but bamboos are common in<br />
much disturbed areas.<br />
The upper tropic<strong>al</strong> rain forest occupies the slope of 600 - 900 m<br />
<strong>al</strong>titude, and are <strong>al</strong>so two stories. The upper story is represented by a great propotion of<br />
oaks and chestnut (Quercus, Lithocarpus and Castanopsis), interspersed with<br />
Magnolia, Michelia, Syzygium, <strong>et</strong>c. The lower story is composed of Antidesma, Aglaia,<br />
Bacaurea, <strong>et</strong>c. P<strong>al</strong>ms are abundant, undergrowth is dense, climbers are few, and<br />
scattered, but epiphyte are abundant. Trees are heavily covered with mosses, fern and<br />
orchids. This is a transition<strong>al</strong> area b<strong>et</strong>ween the hill evergreen or lower mountain forest,<br />
which starts from the elevation of 1,000 m upwards.<br />
1.1.2 Dry or Semi-Evergreen Forest<br />
Gener<strong>al</strong>ly, the dry evergreen forest has a structure similar to the<br />
tropic<strong>al</strong> rain forest but with more xeric climate. The species composition of this type<br />
differs from the tropic<strong>al</strong> rain forest because of different climate and physiography. The<br />
dry evergreen forest is scattered throughout the country <strong>al</strong>ong the depressions on the<br />
peneplain, <strong>al</strong>ong the v<strong>al</strong>leys of low hill range of about 500 m elevation, or <strong>al</strong>ong streams<br />
and rivul<strong>et</strong>s. The rainf<strong>al</strong>l where it occurs is b<strong>et</strong>ween 1,000-2,000 mm. It is difficult to<br />
define this forest type by elevation because it is usu<strong>al</strong>ly found <strong>al</strong>ong the stream banks.<br />
Characteristic tree species are Hopea odorata Roxb., H. ferrea Pierre, Anisoptera spp.,<br />
and Dipterocarpus spp. P<strong>al</strong>m are scattered and belong to the genera of C<strong>al</strong>lamus,<br />
Areca, Livistona, Corypha, and Rhaphis (Siripornnoppakun, 1989).<br />
1.1.3 Hill or Lower Mountain Forest<br />
19
Hill or lower mountain forest is confined above an <strong>al</strong>titude of 1,000<br />
m.; it is scattered <strong>al</strong>l over the country, but mostly found in the northwestern highlands<br />
(Siripornnoppakun, 1989). The atmospheric humidity is very high, which explain the<br />
moss-clad trees; the precipitation is 1,500-2,000 mm annu<strong>al</strong>ly. This forest type is<br />
characterized by Quercus spp., Castanopsis spp., and Lithocarpus spp. Trees and<br />
undergrowth are dense.<br />
1.2 Coniferous Forest<br />
This type of forest is a scattered in sm<strong>al</strong>l pock<strong>et</strong> of the Northwest highlands<br />
and the Khorat Plateau at elevation ranges of 400 to 1,300 m with poor acid soils. The<br />
annu<strong>al</strong> rainf<strong>al</strong>l is about 100-1500 mm. Dominant trees species are Pinus merkusii Jungh<br />
& Devriese, and P. kesiya Royle ex Gordon (Siripornnoppakun, 1989).<br />
1.3 Swamp Forest<br />
Along the depressions in low lying land, the estuaries and mud seashores, a<br />
unique type of veg<strong>et</strong>ation occurs. This type is more or less subjected to periodic<br />
inundation, and is scattered in the w<strong>et</strong> region of the country, where the annu<strong>al</strong> rainf<strong>al</strong>l is<br />
high (exceeding 2000 mm). Owing to the water-logged soil, certain species develop a<br />
characteristic rooting system, bearing pheumatophores, they may up-end above the<br />
ground as knob, or knee-root or prop-roots which sprout from the bas<strong>al</strong> part of the stem<br />
forming tangled masses to anchor it; others <strong>al</strong>so develop buttress.<br />
The forest can be physiographic<strong>al</strong>ly classified into two sub-types, the Fresh<br />
Water Swamp forest and the Mangrove Swamp forest.<br />
1.3.1 Fresh water swamp forest. This type of forest is usu<strong>al</strong>ly found <strong>al</strong>ong<br />
depressions inland. The soil is either <strong>al</strong>luvi<strong>al</strong> or sandy. If it is on <strong>al</strong>luvium the surface is<br />
muddy and fen-like.<br />
20
1.3.2 Mangrove swamp forest. Mangrove swamps are to be found <strong>al</strong>ong<br />
the estuaries of river and the muddy coastlines, where the soil is deep <strong>al</strong>luvium with a<br />
high s<strong>al</strong>ine content. The forest is daily inundated by the tide. In Thailand this type of<br />
forest is very extensive on the West coasty starting from Satui and continuing northwards<br />
to Ranong, and within the Gulf of Thailand occurs from Samut Sakron. In the southwest<br />
to Trat in the southeast.<br />
Species inhabiting this type of forest are semi-xerophytic; they are<br />
occasion<strong>al</strong>ly submerged by seawater, of which they can make no use. The thick leathery<br />
are characteristic, and serve the plants for water storage.<br />
1.4 Beach Forest<br />
On coast<strong>al</strong> sand dunes, rocky seashores and elevated seashore coat<br />
another type of forest occurs; it is gener<strong>al</strong>ly know as Beach forest. This type of forest is<br />
most common <strong>al</strong>ong the East Coast, where the fullest impact of the wind, rain and waves<br />
is felt. Even there, however, only sm<strong>al</strong>l strips or patches of this veg<strong>et</strong>ation type exist. See<br />
the enclosed map.<br />
2. Deciduous Forest<br />
Along the dry belt of the country, where rainf<strong>al</strong>l is lowest (under 1,000 mm), the<br />
climate is more season<strong>al</strong>. The soil is either sandy or gravelly loam; it is som<strong>et</strong>imes<br />
lateristic. The veg<strong>et</strong>ation of these regions is classified as Deciduous, since trees shed<br />
their leaves during the dry season. Trees growing in this type of forest tend to develop<br />
growth or annu<strong>al</strong> rings, a feature not often found in the species of the evergreen forest.<br />
This forest is more or less subjected to ground fire during the dry season.<br />
Deciduous forest can be sub-divided into three main sub-type:<br />
2.1 Mixed Deciduous Forest<br />
21
2.2 Dry Dipterocarp Forest<br />
2.3 Savannah Forest<br />
2.1 Mixed Deciduous Forest<br />
This type of forest is usu<strong>al</strong>ly composed of sever<strong>al</strong> deciduous species in a<br />
mixed association, but in certain loc<strong>al</strong>ities a single species may become predominant;<br />
teak (Tectona grandis) is an example and forest in which this species predominates are<br />
gener<strong>al</strong>ly termed Teak forest for convenience.<br />
Mixed Deciduous forests can <strong>al</strong>so be classified into three sub-types,<br />
based on the terrain and climatic conditions:<br />
2.1.1 Moist Upper Mixed Deciduous<br />
2.1.2 Dry Upper Mixed Deciduous<br />
2.1.3 Lower Mixed Deciduous<br />
2.1.1 Moist Upper Mixed Deciduous Forest. This type of forest occurs<br />
b<strong>et</strong>ween the elevations of 300 - 500 m. and is three storied in profile. The soil bearing<br />
this forest is usu<strong>al</strong>ly loamy, either c<strong>al</strong>careous or granitic. A sm<strong>al</strong>l number of p<strong>al</strong>ms are to<br />
be found scattered in this type of forest. Bamboo are <strong>al</strong>so present.<br />
2.1.2 Dry Upper Mixed Deciduous. Along the ridges at an elevation of<br />
300 - 500 m., the veg<strong>et</strong>ation becomes more open, due to the high rate of evaporation,<br />
exposure and surface erosion and leaching of organic components of the soil. The soil<br />
is either sandy loam or gravel.<br />
The forest is <strong>al</strong>so three storied. Species occurring in the former<br />
type are <strong>al</strong>so present but become rather stunted and crooked. The ground flora is<br />
frequently destroyed by fire. This type of forest, especi<strong>al</strong>ly when regularly disturbed by<br />
22
man will degrade into a bamboo sward, which som<strong>et</strong>imes covers quite an extensive<br />
area.<br />
2.1.3 Lower Mixed Deciduous Forest. This forest type occurs in lowlying<br />
country at an elevation of 50-300 m. in the dry zone, where the soil is colluvi<strong>al</strong> and<br />
either sandy loam or lateritic. As in the former type, the forest is three-storied. The<br />
absence of Teak (Tectona grandis) from the upper story is a distinct characteristic and<br />
serves as a criterion to differentiate this type from the Upper Mixed Deciduous in<br />
Northern Thailand.<br />
2.2 Dry Dipterocarp Forest<br />
On undulating peneplains and ridges where the soil is either sandier<br />
gravelly and subjected to extreme leaching and erosion, the veg<strong>et</strong>ation is markedly<br />
different. The predominant species belong to the family Dipterocarpaceae, hence the<br />
name. The forest is rather open and can be considered as two-storied.<br />
2.3 Savannah Forest<br />
Savannah can be deemed the most extreme from of deciduous type, and<br />
has originated subsequent to burning. It is most frequent in the northeastern region,<br />
where shifting cultivation has been practiced from time immemori<strong>al</strong>. The soil is either<br />
sandy or lateric. Rainf<strong>al</strong>l is relatively low (50-500 mm.). Sm<strong>al</strong>l patches of savannah of<br />
different stages are scattered <strong>al</strong>l over the region. The most extensive ones become vast<br />
grassland, such as Thung Kula Rawng Hai in Sakol Nakhon.<br />
23
1. Study sites<br />
MATERIALS AND METHODS<br />
Materi<strong>al</strong>s<br />
Two sites of forest reserves were selected for the study. Khao Kitchagoot<br />
Nation<strong>al</strong> Park and Khao Soi Dao Wildlife Sanctuary were selected as study sites. Both<br />
sites are in Chanthaburi Province.<br />
1.1 Khao Kitchagoot Nation<strong>al</strong> Park<br />
Khao Kitchagoot Nation<strong>al</strong> Park locates in Amphur Makham, Chanthaburi<br />
Province, Eastern Thailand (latitude 12 49’ N, 102 9’ E). This nation<strong>al</strong> park is part of<br />
eastern forest complex. The topography of this site is a rather steep slope. The elevation<br />
measurement is approximately from 120-1,085 m above mean sea level. The main forest<br />
types are moist evergreen forest (Figure 1) and hill evergreen forest with densely<br />
agricultur<strong>al</strong> areas on the nation<strong>al</strong> park boundary.<br />
The area is influenced mainly by two monsoons, namely the northeast and<br />
southwest monsoons. Winter season occurs during November-February, the summer<br />
season from March-April, and rainy season from May-October. During 1969-1998,<br />
average annu<strong>al</strong> rainf<strong>al</strong>l was 2,865 mm, average relative humidity was 79% and average<br />
temperature was 27.0 C (Wachrinrat <strong>et</strong> <strong>al</strong>., 2001 a).<br />
1.2 Khao Soi Dao Wildlife Sanctuary<br />
Khao Soi Dao Wildlife Sanctuary locates partly in Amphur Pong Num Rorn,<br />
Amphur Makham and Amphur Tha-Mai, Chanthaburi Province, Eastern Thailand (latitude<br />
13 05’ N, 102 10’ E). This wildlife sanctuary is <strong>al</strong>so a part of eastern forest complex.<br />
24
The topography of this site is <strong>al</strong>so a rather steep slope. The elevation measurement is<br />
approximately from 200-1,600 m above mean sea level. The main forest type is dry<br />
evergreen forest (Figure 2) with agricultur<strong>al</strong> areas and fruit orchards <strong>al</strong>ong the wildlife<br />
sanctuary boundary.<br />
The area is <strong>al</strong>so influenced the northeast and southwest monsoons. Winter<br />
season occurs during November-February, the summer season from March-April, and<br />
rainy season from May-October. During 1978-1995, average annu<strong>al</strong> rainf<strong>al</strong>l was 1,517<br />
mm, average relative humidity was 73% and average temperature was 27.0 C<br />
(Wachrinrat <strong>et</strong> <strong>al</strong>., 2001 a).<br />
2. Lay out of Sampling plots<br />
The size of the sampling plots were 100 m x 100 m for each forest type. Sixty<br />
sample plots or 60% of the study area were randomly surveyed in each forest type<br />
(Figure 3). The survey was made 2 time at different season of the year: the cool – dry<br />
season (December-January) and hot – dry season (April); starting from December 1999.<br />
3. Tools and equipment<br />
3.1 Vi<strong>al</strong> 1-2 dram.<br />
3.2 80% <strong>et</strong>hanol.<br />
3.3 Forceps and chicken feathers.<br />
3.4 Knives and axes.<br />
3.5 Sm<strong>al</strong>l hand-spade, shovel, pix-axe.<br />
3.6 Plastic tray.<br />
3.7 Stereo-microscope.<br />
3.8 Identification key of Ahmad (1965), Morimoto (1973) and Tho (1992).<br />
25
1. Survey and sampling m<strong>et</strong>hods.<br />
M<strong>et</strong>hods<br />
1.1 Observations were made on every possible locations where termites would<br />
occur such as tunneling and foraging characteristics: in living trees, branches and stem<br />
or in broken branches or logs on ground.<br />
1.2 Investigations were <strong>al</strong>so made to every common food source of termites<br />
such as tree stumps, wood debris and leaves and litters on ground.<br />
1.3 Observation on nest characteristics was <strong>al</strong>so exercised and the termite<br />
habitats food sources and nest characteristics in each plot of each forest type were<br />
recorded and mapping.<br />
2. Sample collection<br />
2.1 Samples were collected using forceps to collect as many different termite<br />
castes as possible. Soldiers are the main caste used for identification, so it is important<br />
to collect ample individu<strong>al</strong>s when possible. Collecting termites were preserved in vi<strong>al</strong>s<br />
containing 80% <strong>et</strong>hanol and location, date and name of collectors were labeled on<br />
paper and put inside the vi<strong>al</strong>.<br />
2.2 Termite samples were identified using a stereo-microscope based on their<br />
morphologic<strong>al</strong> key index of Ahmad (1965), Morimoto (1973) and Tho (1992).<br />
26
3. Data an<strong>al</strong>ysis<br />
Species diversity of termites in the study area c<strong>al</strong>culated using number of termite<br />
species in a subplot of moist evergreen forest and dry evergreen forest. The Shannon-<br />
Weiner index was used with the following model (Shannon and Weaver, 1949; Pielou,<br />
1975).<br />
s<br />
H’ = -Σp ilog ep i<br />
i=1<br />
Where as: H’ = the v<strong>al</strong>ue of the Shannon-Wiener diversity index.<br />
pi = the proportion of ith species in the whole community.<br />
loge = the natur<strong>al</strong> logarithm of pi. s = the number of species in the community.<br />
Presence of a termite frequency in a subplot was counted as only once.<br />
27
Figure 1 The study site in moist evergreen forest.<br />
Figure 2 The study site in dry evergreen forest.<br />
28
10<br />
11 30 31 50 50 70 71 90 91<br />
9 12 29 32 49 51 69 72 89 92<br />
8 13 28 33 48 52 68 73 88 93<br />
7 14 27 34 47 53 67 74 87 94<br />
6 15 26 35 46 54 66 75 86 95<br />
5 16 25 36 45 55 65 76 85 96<br />
4 17 24 37 44 56 64 77 84 97<br />
3 18 23 38 43 57 63 78 83 98<br />
2 19 22 39 42 58 62 79 82 99<br />
1 20 21 40 41 60 61 80 81<br />
100<br />
= Sample plots were randomly surveyed<br />
Figure 3 Model of sampling plots in moist evergreen forest and<br />
dry evergreen forest.<br />
29
RESULTS AND DISCUSSION<br />
Three hundred and forty five samples of termite were collected from moist<br />
evergreen forest (MEF) at Khao Kitchagoot Nation<strong>al</strong> Park and dry evergreen forest (DEF)<br />
at Khao Soi Dao Wildlife Sanctuary. Their morphologic<strong>al</strong> characteristics were<br />
investigated base on the systematic keys of Ahmad (1965), Morimoto (1973) and Tho<br />
(1992). Results reve<strong>al</strong>ed that tot<strong>al</strong>ly 42 species belong to 20 genera, 8 subfamilies, 3<br />
families were collected from these areas (Figure 4). The d<strong>et</strong>ail use in the identification<br />
and the characteristics of each subfamily are described as follows:<br />
No. of termites<br />
14<br />
12<br />
10<br />
8<br />
6<br />
4<br />
2<br />
0<br />
4<br />
<br />
<br />
K<strong>al</strong>otermitinae<br />
2<br />
<br />
<br />
<br />
1<br />
Rhinotermitinae<br />
<br />
<br />
<br />
5<br />
<br />
<br />
<br />
<br />
1 1<br />
<br />
Coptotermitinae<br />
<br />
<br />
4<br />
<br />
<br />
Amitermitinae<br />
<br />
<br />
<br />
<br />
No. of species<br />
<br />
2<br />
9<br />
<br />
<br />
Macrotermitinae<br />
Subfamily<br />
5<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
13<br />
<br />
<br />
<br />
<br />
<br />
<br />
Nasutitermitinae<br />
3<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
No. of genera<br />
<br />
4<br />
Termitinae<br />
4<br />
<br />
<br />
<br />
<br />
<br />
<br />
2<br />
Apicotermitinae<br />
2<br />
<br />
<br />
<br />
Figure 4 Species diversity of termite groups in the study sites (moist evergreen<br />
forest and dry evergreen forest).<br />
30
1. Subfamily K<strong>al</strong>otermitinae<br />
This is the wood-dwellers that confined throughout of their life in wood which is<br />
usu<strong>al</strong>ly above ground. The wood may be parts of living or dead tree of which the<br />
swarming pair enters to start a new colony (Roonw<strong>al</strong>, 1970). Two genera and four<br />
species of this subfamily were observed.<br />
Glyptotermes and Cryptotermes are among the majority species of this<br />
subfamily. Glyptotermes is a damp-wood termites which require wood with a constant<br />
high amount of moisture and usu<strong>al</strong>ly found in dead or decaying logs or tree stumps in<br />
moist, cool regions while Cryptotermes is a dry-wood termite which able to flourish in<br />
dry, sound, seasoned wood (Roonw<strong>al</strong>, 1970). Chhotani (1970) reported that<br />
Cryptotermes is a dominant genus of K<strong>al</strong>otermitidae that is a typic<strong>al</strong> dry wood termite, it<br />
is widely distributed in tropic<strong>al</strong> region but tend to be confined to forests <strong>al</strong>ong coast<strong>al</strong><br />
and islands.<br />
2. Subfamily Rhinotermitinae<br />
This subfamily belongs to a relatively primitive family with a worldwide<br />
distribution. It is speculated as being orient<strong>al</strong> in origin<strong>al</strong> (Emerson, 1955; Krishna, 1970).<br />
Roonw<strong>al</strong> (1970) described that this subfamily attacks rotten wood, weathered timber,<br />
dead branches and wounds of many trees including p<strong>al</strong>ms. This subfamily usu<strong>al</strong>ly has<br />
broad earth-covered g<strong>al</strong>leries with irregular surface run on tree trunks and on wood work<br />
in houses. Nests are constructed inside stumps of old trees, in wounds in forest trees at<br />
ground level and in cavities in beams of wood. One genus and one species of this<br />
subfamily were observed.<br />
3. Subfamily Coptotermitinae<br />
31
Roonw<strong>al</strong> (1970) described that every species in this subfamily attacks wood and<br />
some of them are serious pests of woodwork in house. Nesting is gener<strong>al</strong>ly underground<br />
or inside the trunks of trees, in logs and stumps, b<strong>et</strong>ween wooden boards, in house and<br />
in railway carriages. One genus and five species of this subfamily were observed.<br />
Coptotermes gestroi Wasmann, the most serious pest causing damage in<br />
buildings or wooden constructions in Thailand (Sornnuwat, 1996). Roonw<strong>al</strong> (1970)<br />
reported that this species is a serious pest of living rubber trees in Southeast Asia and<br />
attacking bark and other portions, including the heartwood which often killing the plant.<br />
It <strong>al</strong>so attacks a few other species of trees. He <strong>al</strong>so described that this species reach its<br />
food it makes underground tunnels and, if necessary, covered runways of digested<br />
wood and soil particles are made above ground.<br />
4. Subfamily Macrotermitinae<br />
This is the most important termite groups which play an important role in the<br />
process of litter decomposition with higher weight specific consumption rates than other<br />
termites (Collins, 1981; 1983). They feed on dead wood and leaves, often in relatively<br />
fresh condition, by the action of their mutu<strong>al</strong>istic fungi. These termites use only soil and<br />
s<strong>al</strong>iva to build their mound (Noirot, 1970) and play an important role in the turnover of<br />
organic matter and the maintenance or improvement of soil fertility (Krishna and<br />
Weesner, 1969; 1970). Five genera and nine species of this subfamily were observed.<br />
5. Subfamily Amitermitinae<br />
Roonw<strong>al</strong> (1970) described that this subfamily is characterized by soldiers with a<br />
subrectangular head gener<strong>al</strong>ly having saber-shaped mandibles with a single fairly<br />
prominent tooth or with the margin more or less serrated. Two genera and four species<br />
of this subfamily were observed.<br />
32
Microcerotermes is the main genus found widely distributed over <strong>al</strong>l the<br />
zoogeographic<strong>al</strong> regions except the Nearctic, this genus is wood-eating species and<br />
carton-nest builders; construct cellular nest of carton (a substance made of a mixture of<br />
termite fec<strong>al</strong> matter, woody fragments, and som<strong>et</strong>ime earth). The carton nests may be<br />
arbore<strong>al</strong> or partly or wholly buried in the earth (Roonw<strong>al</strong>, 1970).<br />
Globitermes sulphureus (Haviland) was another termite in Amitermitinae widely<br />
distributed in survey sites. Their dome shape mound <strong>al</strong>ways found above ground and<br />
usu<strong>al</strong>ly covered extern<strong>al</strong>ly with thin layer of earth (Roonw<strong>al</strong>, 1970). Tho (1992) reported<br />
their habitats to be widespread throughout the lowlands of M<strong>al</strong>aysia, especi<strong>al</strong>ly on the<br />
west coast. He described that this species was very common in rubber, coconut, oil<br />
p<strong>al</strong>m and sugar cane plantation.<br />
6. Subfamily Nasutitermitinae<br />
Higashi and Abe (1997) explained that the Nasutitermitinae, which contain more<br />
than 500 species, nest in various sites such as in dead wood, on tree trunks, on the<br />
ground, and in soil. They consume various dead plant materi<strong>al</strong>s such as dead wood,<br />
dead grasses, soil, dung, and <strong>al</strong>so lichen. Nasutitermitinae commonly built their nests on<br />
very high trees over 5-10 m<strong>et</strong>ers (arbore<strong>al</strong> nest) and feed on semi-decomposed leaf and<br />
twigs in contact with soil or dead branches. Three genera and 13 species of this<br />
subfamily were observed.<br />
Hospit<strong>al</strong>itermes is strictly orient<strong>al</strong> distribution and <strong>al</strong>l the known species are free<br />
foraging in their foraging behavior (Tho, 1992). Their nests are found in the forks of trees<br />
and mudlike structure hang on their sides. The termite goes out foraging in marching<br />
expedition. The food consists mainly of lichens and <strong>al</strong>so of fungi and <strong>al</strong>gae. The termites<br />
undertake long foraging expeditions to distant place. After collecting the food in the<br />
mouth, the workers r<strong>et</strong>urn home som<strong>et</strong>ime shortly before dawn, and in later stages the<br />
procession consists of an outgoing and incoming stream side by side (Roonw<strong>al</strong>, 1970).<br />
33
7. Subfamily Apicotermitinae<br />
Noirot (1970) described that <strong>al</strong>l termites in Apicotermitinae are humivores and<br />
construct their nests in the soil. Noirot and Noirot-Timothee (1969) defined that humivore<br />
is the eaters of humus having a regime which varies more from the primitive di<strong>et</strong>. It is<br />
unknow just how these termites utilize the humus, but they probably use the more or less<br />
decomposed veg<strong>et</strong>able debris, which it contains. In addition this subfamily are<br />
soldierless and soil feeders that form diffuse underground nest in the soil (Roonw<strong>al</strong>,<br />
1970; Eggl<strong>et</strong>on, 2000). Two genera and two species of this subfamily were observed.<br />
8. Subfamily Termitinae<br />
Similar to the subfamily Apicotermitinae discussed previously, Termitinae are<br />
humivores feeding on substrates such as dung and plant litter in various stages of<br />
decomposition (Lee and Wood, 1971). They are usu<strong>al</strong>ly classified as subterranean<br />
termite and their nests are found situated underground excepted some genus such as<br />
Termes cosmis Haviland which their nests are either partly buried in the ground or made<br />
on branches of trees close to the ground or in the hollows of trees (Ahmad, 1965;<br />
Roonw<strong>al</strong>, 1970). Four genera and four species of this subfamily were observed.<br />
Specification of Results from Collected Specimens<br />
Termites from the result obtained from the identification reported earlier were<br />
further specified into 3 main groups as followed:-<br />
1. Morphologic<strong>al</strong> characteristic.<br />
2. Food habitat.<br />
3. Nest habitat.<br />
1. Termite genera and species classified by their morphologic<strong>al</strong> characteristics<br />
34
1.1 Moist evergreen forest<br />
Thirty-seven species of 19 genera and 8 subfamilies were recorded from<br />
moist evergreen forest (Table1 and Figure 5):<br />
1.1.1 K<strong>al</strong>otermitinae: Three und<strong>et</strong>ermined species of genus<br />
Glyptotermes and 1 species of Cryptotermes.<br />
1.1.2 Rhinotermitinae: One species of Schedorhinotermes<br />
(Schedorhinotermes medioobscurus (Holmgren)).<br />
1.1.3 Coptotermitinae: Three species of Coptotermes: Coptotermes<br />
gestroi Wasmann, C. havilandi Holmgren and C. premrasmii Ahmad.<br />
1.1.4 Amitermitinae: Two species of Microcerotermes and 1 species of<br />
Globitermes (Globitermes sulphureus (Haviland)).<br />
1.1.5 Macrotermitinae: Four species of Odontotermes, 2 species of<br />
Macrotermes and 1 species each of Ancistrotermes, Microtermes and Hypotermes.<br />
1.1.6 Nasutitermitinae: Five species of Nasutitermes, 4 species of<br />
Bulbitermes and 3 species of Hospit<strong>al</strong>itermes.<br />
1.1.7 Termitinae: One species each of genus Termes, Mirocapritermes<br />
and Angulitermes were found in moist evergreen forest. Interestingly, Angulitermes was<br />
report as the newly recorded termite of Thailand (Vongk<strong>al</strong>uang <strong>et</strong> <strong>al</strong>., 2001) (Figure 7).<br />
1.1.8 Apicotermitinae: Two species of genus Speculitermes and<br />
Euhamitermes.<br />
1.2 Dry evergreen forest<br />
Twenty-seven species of 15 genera and 7 subfamilies were recorded from<br />
dry evergreen forest (Table1 and Figure 5):<br />
1.1.1 K<strong>al</strong>otermitinae: Two species each of Cryptotermes and<br />
Glyptotermes.<br />
35
1.1.2 Rhinotermitinae: One species of Schedorhinotermes<br />
(Schedorhinotermes medioobscurus (Holmgren)).<br />
1.1.3 Coptotermitinae: Three species of Coptotermes: Coptotermes<br />
gestroi Wasmann, C. curvignathus Holmgren and one und<strong>et</strong>ermined species of<br />
Coptotermes.<br />
1.1.4 Amitermitinae: Two species of Microcerotermes and 1 species of<br />
Globitermes (Globitermes sulphureus (Haviland)).<br />
1.1.5 Macrotermitinae: Two species of Odontotermes and 1 species<br />
each of genus Macrotermes, Ancistrotermes, Microtermes and Hypotermes.<br />
1.1.6 Termitinae: One species each of genus Termes and<br />
Pericapritermes.<br />
1.1.7 Nasutitermitinae: Six species of Nasutitermes and 4 species of<br />
Bulbitermes.<br />
36
TABLE 1 Termite genera and species recorded in two different forest types: moist<br />
evergreen forest and dry evergreen forest. MEF = moist evergreen forest,<br />
DEF = dry evergreen forest, F = number of family, SF = number of<br />
subfamily, G = number of genera, SP = number of species.<br />
Termite Genus and Species Forest types<br />
(3F, 8SF, 20G, 42SP) MEF DEF<br />
1. F. K<strong>al</strong>otermitidae (2G, 4SP)<br />
1.1 SF. K<strong>al</strong>otermitinae<br />
1. Cryptotermes thailandis Ahmad + +<br />
2. Glyptotermes sp.1 + -<br />
3. Glyptotermes sp.2 + +<br />
4. Glyptotermes sp.3 + -<br />
2. F Rhinotermitidae (2G, 6SP)<br />
2.1 SF Rhinotermitinae (1G, 1SP)<br />
5. Schedorhinotermes medioobscurus (Holmgren) + +<br />
2.2 SF Coptotermitinae (1G, 5SP)<br />
6. Coptotermes sp.1 - +<br />
7. Coptotermes gestroi Wasmann + +<br />
8. Coptotermes havilandi Holmgren + -<br />
9. Coptotermes premrasmii Ahmad + -<br />
10. Coptotermes curvignathus Holmgren - +<br />
3. F Termitidae (16G, 32SP)<br />
3.1 SF Macrotermitinae (5G, 9SP)<br />
11. Macrotermes annand<strong>al</strong>ei (Silvestri) + +<br />
12. Macrotermes maesodensis Ahmad + -<br />
13. Microtermes obesi Holmgren + +<br />
14. Ancistrotermes pakestanicus (Silvestri) + +<br />
15. Hypotermes makhamensis Ahmad + +<br />
16. Odontotermes sp.1 + -<br />
17. Odontotermes proformosanus Ahmad + +<br />
18. Odontotermes formosanus (Shiraki) + +<br />
19. Odontotermes oblongathus Holmgren + -<br />
3.2 SF Amitermitinae (2G,4SP)<br />
20. Microcerotermes crassus Snyder + +<br />
21. Microcerotermes distans (Haviland) + +<br />
22. Microcerotermes annand<strong>al</strong>ei Silvestri + -<br />
23. Globitermes sulphureus (Haviland) + +<br />
3.3 SF Apicotermitinae (2G,2SP)<br />
24. Euhamitermes sp. + -<br />
25. Speculitermes sp. + -<br />
37
TABLE 1 (cont.) Termite genera and species recorded in two different forest types:<br />
moist evergreen forest and dry evergreen forest. MEF = moist<br />
evergreen forest, DEF = dry evergreen forest, F = number of family,<br />
SF = number of subfamily, G = number of genera, SP = number of<br />
species.<br />
Termite Genus and Species Forest types<br />
(3F, 8SF, 20G, 42SP) MEF DEF<br />
3.4 SF Termitinae (4G, 4SP)<br />
26. Termes cosmis Haviland + +<br />
27. Pericapritermes sp.C - +<br />
28. Mirocapritermes concaveus Ahmad + -<br />
29. Angulitermes sp. + -<br />
3.5 SF Nasutitermitinae (3G, 13SP)<br />
30. Nasutitermes johoricus (John) + +<br />
31. Nasutitermes matangensiformis (Holmgren) - +<br />
32. Nasutitermes sp.1 + +<br />
33. Nasutitermes sp.3 - +<br />
34. Nasutitermes sp.4 - +<br />
35. Nasutitermes sp.5 + +<br />
36. Bulbitermes parapusillus Ahmad + +<br />
37. Bulbitermes laticeph<strong>al</strong>us Ahmad + +<br />
38. Bulbitermes sp.1 + +<br />
39. Bulbitermes sp.2 + +<br />
40. Hospit<strong>al</strong>itermes ataramensis Prashad and Sen-Sarma + -<br />
41. Hospit<strong>al</strong>itermes sp.3 + -<br />
42. Hospit<strong>al</strong>itermes sp.4 + -<br />
38
No. of species and genera<br />
Results of survey as shown in Table 1 and Figure 5 reve<strong>al</strong>ed that moist<br />
evergreen forest showed the higher number of termite species than dry evergreen forest<br />
(having tot<strong>al</strong>ly 36 species of 19 genera, 3 families to 27 species of 15 genera, 3<br />
families).<br />
No. of species<br />
40<br />
30<br />
20<br />
10<br />
0<br />
14<br />
12<br />
10<br />
8<br />
6<br />
4<br />
2<br />
0<br />
4<br />
K<strong>al</strong>otermitinae<br />
2<br />
<br />
<br />
<br />
<br />
<br />
37<br />
<br />
<br />
<br />
<br />
1<br />
Rhinotermitinae<br />
1<br />
<br />
<br />
<br />
3<br />
Coptotermitinae<br />
19<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
Figure 5 Species diversity of termite groups in moist evergreen forest (MEF)<br />
and dry evergreen forest (DEF).<br />
3<br />
<br />
<br />
<br />
<br />
<br />
Nasutitermitinae<br />
MEF<br />
12<br />
10<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
Macrotermitinae<br />
Subfamily<br />
9<br />
6<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
MEF DEF<br />
Forest types<br />
<br />
<br />
No. of Species<br />
Amitermitinae<br />
3 3 3<br />
<br />
<br />
<br />
<br />
<br />
DEF<br />
Termitinae<br />
2<br />
<br />
<br />
<br />
<br />
<br />
2<br />
<br />
<br />
Apicotermitinae<br />
Figure 6 Species of termites in moist evergreen forest (MEF) and dry evergreen<br />
forest (DEF).<br />
27<br />
<br />
<br />
No. of Genera<br />
15<br />
0<br />
39
From the results which reve<strong>al</strong>ed that Glyptotermes (subfamily K<strong>al</strong>otermitinae)<br />
was found in higher number of species in moist evergreen forest than dry evergreen<br />
forest (Table1 and Figure 6) might be due to the fact that moist evergreen forest having<br />
higher average rainf<strong>al</strong>l and relative humidity than dry evergreen forest (Wachrinrat <strong>et</strong> <strong>al</strong>.,<br />
2001 a) and may be more suitable for Glyptotermes, which is the damp-wood termite, to<br />
establish their colony.<br />
The reason for the presence of Schedorhinotermes medioobscurus (Holmgren)<br />
(subfamily Rhinotermitinae) in both forest types might be because these forest types<br />
have ample rotten wood and litterf<strong>al</strong>l including branches and stems (Wachrinrat <strong>et</strong> <strong>al</strong>.,<br />
2001 b) for this termite to forage.<br />
There are no difference in the number of species of subfamily Coptotermitinae in<br />
moist evergreen forest and dry evergreen forest (Table 1 and Figure 6). Therefore it was<br />
assumed that these 2 forest types probably contain sufficient amount of trees and wood<br />
for these termites to forage.<br />
Two species of Apicotermitinae were found specific<strong>al</strong>ly in moist evergreen forest<br />
(Table 1 and Figure 6). Sawasdee <strong>et</strong> <strong>al</strong>. (2001) reported that the average depth of soil<br />
litter surface thickness in moist evergreen forest was 7.40 cm that which thicker than dry<br />
evergreen forest (5.68 cm). The average water content at the depth 0-15 cm in moist<br />
evergreen forest and dry evergreen forest were 0.270 and 0.173 respectively.<br />
Percentages of organic matter were 5.55% for moist evergreen forest and 3.99% for dry<br />
evergreen forest. The information above lead to the speculation that moist evergreen<br />
forest probably has appropriate soil organic matter and soil moisture for Apicotermitinae<br />
to survive than dry evergreen forest.<br />
Similar to the subfamily Apicotermitinae. Number of species of Termitinae was<br />
found in higher degree in moist evergreen forest than in dry evergreen forest. Nests of<br />
40
this subfamily are mostly found underground excepted Termes cosmis Haviland which<br />
their nests are either partly buried in the ground or made on branches of trees close to<br />
the ground or in the hollows of trees (Ahmad, 1965; Roonw<strong>al</strong>, 1970).<br />
For subfamily Nasutitermitinae, three species of Hospit<strong>al</strong>itermes were found only<br />
in moist evergreen forest it might be due to the food of this termite group consists mainly<br />
of lichen (Roonw<strong>al</strong>, 1970). From these survey we can found this termites carry lichen in<br />
their mouth particularly in moist evergreen forest. Abe (1979) reported that 2 species of<br />
Hospit<strong>al</strong>itermes in Pasoh Forest were observed very commonly to collect scraping of<br />
lichen usu<strong>al</strong>ly on living tree trunks and som<strong>et</strong>imes on f<strong>al</strong>len trees. Wachrinrat <strong>et</strong> <strong>al</strong>. (2001<br />
a) reported that moist evergreen forest has highest moisture that might be suitable for<br />
Hospit<strong>al</strong>itermes to live.<br />
41
0.5 mm.<br />
A<br />
Figure 7 Soldier and worker of Angulitermes, newly recorded of termite list<br />
of Thailand. A: Whole body of soldier from above; B: Whole body<br />
of worker from above; C: Whole body of soldier from side.<br />
C<br />
B<br />
42
2. Termite group classified by their food habitat.<br />
Classifications of this group gener<strong>al</strong>ly relied on observations of the location of<br />
foraging g<strong>al</strong>leries on ground, under leaves, tree, branches, log or dead stumps and nest<br />
in soil or above ground, <strong>al</strong>so organic materi<strong>al</strong> carried to the nest for storage which<br />
som<strong>et</strong>imes reflex in the color of worker abdomen. Termites collected in moist evergreen<br />
forest and dry evergreen forest were classified into different groups based on food<br />
habitat (Table 2) as follows:<br />
2.1 Wood feeders<br />
2.2 Wood and leaf feeders<br />
2.3 Soil or humus feeders<br />
2.4 Lichen feeders<br />
43
Table 2 Termite genera and species recorded in moist evergreen forest and dry<br />
evergreen forest, eastern Thailand classified by their food habitat. F =<br />
number of family, SF = number of subfamily, G = number of genera, SP =<br />
number of species, W = Wood, W&L = Wood and leaf, S/H = Soil or humus, L<br />
= Lichen.<br />
Termite Genus and Species Food habitat<br />
(3F, 8SF, 20G, 42SP) W W&L S/H L<br />
1. F. K<strong>al</strong>otermitidae (2G, 4SP)<br />
1.1 SF. K<strong>al</strong>otermitinae<br />
1. Cryptotermes thailandis Ahmad X<br />
2. Glyptotermes sp.1 X<br />
3. Glyptotermes sp.2 X<br />
4. Glyptotermes sp.3 X<br />
2. F Rhinotermitidae (2G, 6SP)<br />
2.1 SF Rhinotermitinae (1G, 1SP)<br />
5. Schedorhinotermes medioobscurus (Holmgren) X<br />
2.2 SF Coptotermitinae (1G, 5SP)<br />
6. Coptotermes sp.1 X<br />
7. Coptotermes gestroi Wasmann X<br />
8. Coptotermes havilandi Holmgren X<br />
9. Coptotermes premrasmii Ahmad X<br />
10. Coptotermes curvignathus Holmgren X<br />
3. F Termitidae (16G, 32SP)<br />
3.1 SF Macrotermitinae (5G, 9SP)<br />
11. Macrotermes annand<strong>al</strong>ei (Silvestri) X<br />
12. Macrotermes maesodensis Ahmad X<br />
13. Microtermes obesi Holmgren X<br />
14. Ancistrotermes pakestanicus (Silvestri) X<br />
15. Hypotermes makhamensis Ahmad X<br />
16. Odontotermes sp.1 X<br />
17. Odontotermes proformosanus Ahmad X<br />
18. Odontotermes formosanus (Shiraki) X<br />
19. Odontotermes oblongathus Holmgren X<br />
3.2 SF Amitermitinae (2G,4SP)<br />
20. Microcerotermes crassus Snyder X<br />
21. Microcerotermes distans (Haviland) X<br />
22. Microcerotermes annand<strong>al</strong>ei Silvestri X<br />
23. Globitermes sulphureus (Haviland) X<br />
3.3 SF Apicotermitinae (2G,2SP)<br />
24. Euhamitermes sp. X<br />
25. Speculitermes sp. X<br />
44
Table 2 (cont.) Termite genera and species recorded in moist evergreen forest and dry<br />
evergreen forest, eastern Thailand classified by their food habitat. F =<br />
number of family, SF = number of subfamily, G = number of genera, SP<br />
= number of species, W = Wood, W&L = Wood and leaf, S/H = Soil or<br />
humus, L = Lichen.<br />
Termite Genus and Species Food habitat<br />
(3F, 8SF, 20G, 42SP) W W&L S/H L<br />
3.4 SF Termitinae (4G, 4SP)<br />
26. Termes cosmis Haviland X<br />
27. Pericapritermes sp.C X<br />
28. Mirocapritermes concaveus Ahmad X<br />
29. Angulitermes sp. X<br />
3.5 SF Nasutitermitinae (3G, 13SP)<br />
30. Nasutitermes johoricus (John) X<br />
31. Nasutitermes matangensiformis (Holmgren) X<br />
32. Nasutitermes sp.1 X<br />
33. Nasutitermes sp.3 X<br />
34. Nasutitermes sp.4 X<br />
35. Nasutitermes sp.5 X<br />
36. Bulbitermes parapusillus Ahmad X<br />
37. Bulbitermes laticeph<strong>al</strong>us Ahmad X<br />
38. Bulbitermes sp.1 X<br />
39. Bulbitermes sp.2 X<br />
40. Hospit<strong>al</strong>itermes ataramensis Prashad and Sen-Sarma X<br />
41. Hospit<strong>al</strong>itermes sp.3 X<br />
42. Hospit<strong>al</strong>itermes sp.4 X<br />
2.1 Wood feeders<br />
Termite in this group feeding on wood and woody litter, including dead<br />
branches still attached to trees (Bignell and Eggl<strong>et</strong>on, 2000) (Figure 9).<br />
In moist evergreen forest 11 species of termites were wood feeding<br />
termites. There are 4 species belong to subfamily K<strong>al</strong>otermitinae, 1 species of subfamily<br />
Rhinotermitidae, 3 species of subfamily Coptotermitinae and 3 species belong to<br />
subfamily Amitermitinae.<br />
45
Nine species of termites in dry evergreen forest were wood feeding<br />
termite. There are 2 species of subfamily K<strong>al</strong>otermitinae, 1 species of subfamily<br />
Rhinotermitinae, 3 species of subfamily Coptotermitinae and 3 species of subfamily<br />
Amitermitinae.<br />
2.2 Wood and leaf feeders<br />
This feeding group forage on leaves and sm<strong>al</strong>l woody items and often<br />
taken back and stored their food temporarily in the nest (Bignell and Eggl<strong>et</strong>on, 2000)<br />
(Figure 10).<br />
Eighteen species of termites found in moist evergreen forest were wood<br />
and leaf feeders. Nine species of this group from this forest type belong to subfamily<br />
Nasutitermitinae and nine species belong to subfamily Macrotermitinae.<br />
For dry evergreen forest, 16 species of termites were wood and leaf<br />
feeders. Ten species from this forest type belong to subfamily Nasutitermitinae and six<br />
species were Macrotermitinae.<br />
2.3 Soil or humus feeders<br />
These termites are humivores and live mostly in soil or subterranean nest<br />
except Termes cosmis Haviland that built their nest on the ground, which is usu<strong>al</strong>ly,<br />
c<strong>al</strong>led epige<strong>al</strong> nest (Figure 11).<br />
In moist evergreen forest, five species of termites were soil or humus<br />
feeders. Three species from this forest type belong to subfamily Termitinae and 2<br />
species belong to Apicotermitinae.<br />
46
Two species of termites in dry evergreen forest belonging to subfamily<br />
Termitinae were soil or humus feeding termites.<br />
2.4 Lichen feeders<br />
Termite in this feeding group were observed very commonly to collect<br />
scraping of lichen usu<strong>al</strong>ly on living tree trunks and som<strong>et</strong>imes on f<strong>al</strong>len trees (Abe,<br />
1979) (Figure 12).<br />
Termites of this feeding group were found only in moist evergreen forest<br />
which 3 species of genus Hospit<strong>al</strong>itermes belong to subfamily Nasutitermitinae.<br />
No. of species<br />
20<br />
15<br />
10<br />
5<br />
0<br />
11<br />
9<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
18<br />
16<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
Wood Wood&leaves<br />
Food habitat<br />
Soil/humus Lichen<br />
MEF<br />
5<br />
DEF<br />
Figure 8 Termite species in differing food habitat in moist evergreen forest (MEF)<br />
and dry evergreen forest (DEF).<br />
2<br />
3<br />
0<br />
47
Classification of feeding groups as shown in Figure 8 reve<strong>al</strong>ed that moist<br />
evergreen forest showed the higher number of wood feeding group than dry evergreen<br />
forest. Wachrinrat <strong>et</strong> <strong>al</strong>. (2001 a) reported that tot<strong>al</strong> aboveground biomass in moist<br />
evergreen forest was 434.732 ton/ha which was higher than in dry evergreen forest<br />
(340.558 ton/ha). Therefore this might be the reason that moist evergreen forest have a<br />
highest number of species of wood feeding termites.<br />
The number of wood and leaf feeding termites found in moist evergreen forest<br />
were not much different from that of dry evergreen forest. This might due to the amount<br />
of tot<strong>al</strong> litterf<strong>al</strong>l in dry evergreen forest and moist evergreen forest that is the main supply<br />
of food for this group was nearly equ<strong>al</strong> in both forest types (Wachrinrat <strong>et</strong> <strong>al</strong>., 2001 b).<br />
The results which reve<strong>al</strong>ed that soil or humus feeding termites were found in<br />
higher number of species in moist evergreen forest than in dry evergreen forest might<br />
be due to the fact that moist evergreen forest having higher organic matter and soil<br />
moisture than dry evergreen forest (Sawasdee <strong>et</strong> <strong>al</strong>., 2001) and may be suitable for soil<br />
or humus feeders to live.<br />
Lichen feeding termites were found only in moist evergreen forest. Trees in moist<br />
evergreen forest are covered with mosses, ferns and orchids (Siripronnoppakun, 1989)<br />
and have higher moisture (Wachrinrat <strong>et</strong> <strong>al</strong>., 2001 a) that might be suitable for this<br />
feeding group to live.<br />
48
Figure 9 Wood feeding termites. A: Cryptotermes; B: Microcerotermes.<br />
A<br />
B<br />
49
Figure 10 Wood and leaf feeding termites. A: Hypotermes; B: Macrotermes.<br />
A<br />
B<br />
50
Figure 11 Soil or humus feeding termites. A: Termes; B: Dicuspiditermes.<br />
A<br />
B<br />
51
Figure 12 Lichen feeding termites: Hospit<strong>al</strong>itermes.<br />
A<br />
52
3. Termite group classification by nest habitat.<br />
All termite species have identifiable nests where colony members are housed<br />
and protected. According to their mode of living, dwell in a vari<strong>et</strong>y of habitats, which are<br />
mostly shut out from light, i.e., in soil, inside wood, in arbore<strong>al</strong> nests, <strong>et</strong>c. Termites<br />
collected from this study were classified into different groups based on their nest habitat<br />
(Table 3) as follows:<br />
3.1 Wood-dwellers<br />
3.2 Ground-dwellers<br />
53
Table 3 Termite genera and species recorded in moist evergreen forest and dry<br />
evergreen forest, eastern Thailand classified by their nest habitat. F =<br />
number of family, SF = number of subfamily, G = number of genera, SP =<br />
number of species, IW = Nest in wood, A = Arbore<strong>al</strong> nest, E = Epige<strong>al</strong> nest,<br />
S = Subterranean nest.<br />
Termite Genus and Species Nest habitat<br />
(3F, 8SF, 20G, 42SP) IW A E S<br />
1. F. K<strong>al</strong>otermitidae (2G, 4SP)<br />
1.1 SF. K<strong>al</strong>otermitinae<br />
1. Cryptotermes thailandis Ahmad X<br />
2. Glyptotermes sp.1 X<br />
3. Glyptotermes sp.2 X<br />
4. Glyptotermes sp.3 X<br />
2. F Rhinotermitidae (2G, 6SP)<br />
2.1 SF Rhinotermitinae (1G, 1SP)<br />
5. Schedorhinotermes medioobscurus (Holmgren) X<br />
2.2 SF Coptotermitinae (1G, 5SP)<br />
6. Coptotermes sp.1 X<br />
7. Coptotermes gestroi Wasmann X<br />
8. Coptotermes havilandi Holmgren X<br />
9. Coptotermes premrasmii Ahmad X<br />
10. Coptotermes curvignathus Holmgren X<br />
3. F Termitidae (16G, 32SP)<br />
3.1 SF Macrotermitinae (5G, 9SP)<br />
11. Macrotermes annand<strong>al</strong>ei (Silvestri) X<br />
12. Macrotermes maesodensis Ahmad X<br />
13. Microtermes obesi Holmgren X<br />
14. Ancistrotermes pakestanicus (Silvestri) X<br />
15. Hypotermes makhamensis Ahmad X<br />
16. Odontotermes sp.1 X<br />
17. Odontotermes proformosanus Ahmad X<br />
18. Odontotermes formosanus (Shiraki) X<br />
19. Odontotermes oblongathus Holmgren X<br />
3.2 SF Amitermitinae (2G,4SP)<br />
20. Microcerotermes crassus Snyder X X<br />
21. Microcerotermes distans (Haviland) X<br />
22. Microcerotermes annand<strong>al</strong>ei Silvestri X<br />
23. Globitermes sulphureus (Haviland) X<br />
3.3 SF Apicotermitinae (2G,2SP)<br />
24. Euhamitermes sp. X<br />
25. Speculitermes sp. X<br />
54
Table 3 (cont.) Termite genera and species recorded in moist evergreen forest and dry<br />
evergreen forest, eastern Thailand classified by their nest habitat. F =<br />
number of family, SF = number of subfamily, G = number of genera, SP<br />
= number of species, IW = Nest in wood, A = Arbore<strong>al</strong> nest, E =<br />
Epige<strong>al</strong> nest, S = Subterranean nest.<br />
Termite Genus and Species Nest habitat<br />
(3F, 8SF, 20G, 42SP) IW A E S<br />
3.4 SF Termitinae (4G, 4SP)<br />
26. Termes cosmis Haviland X X<br />
27. Pericapritermes sp.C X<br />
28. Mirocapritermes concaveus Ahmad X<br />
29. Angulitermes sp. X<br />
3.5 SF Nasutitermitinae (3G, 13SP)<br />
30. Nasutitermes johoricus (John) X<br />
31. Nasutitermes matangensiformis (Holmgren) X<br />
32. Nasutitermes sp.1 X<br />
33. Nasutitermes sp.3 X<br />
34. Nasutitermes sp.4 X<br />
35. Nasutitermes sp.5 X<br />
36. Bulbitermes parapusillus Ahmad X<br />
37. Bulbitermes laticeph<strong>al</strong>us Ahmad X<br />
38. Bulbitermes sp.1 X<br />
39. Bulbitermes sp.2 X<br />
40. Hospit<strong>al</strong>itermes ataramensis Prashad and Sen-Sarma X<br />
41. Hospit<strong>al</strong>itermes sp.3 X<br />
42. Hospit<strong>al</strong>itermes sp.4 X<br />
3.1 Wood-dwellers<br />
This group was described by Roonw<strong>al</strong> (1970) to have their nest confined<br />
throughout their lives wood which is usu<strong>al</strong>ly above ground. Only subfamily<br />
K<strong>al</strong>otermitinae was found in present study (Figure 14).<br />
Four species of termite in moist evergreen forest belong to subfamily<br />
K<strong>al</strong>otermitinae were wood-dwellers.<br />
55
forest.<br />
Two species of subfamily K<strong>al</strong>otermitinae were found in dry evergreen<br />
3.2 Ground-dwellers<br />
The colony of ground dwelling termites <strong>al</strong>ways exists at least partly in the<br />
ground; the colonized pair enters the earth or wood lying in the earth. The ground<br />
dwellers can be further classified into 3 subgroups.<br />
3.2.1 Subterranean termites<br />
3.2.2 Arbore<strong>al</strong> nest<br />
3.2.3 Epige<strong>al</strong> nest<br />
3.2.1 Subterranean termites<br />
This termite group lives underground, but build covered runways to<br />
reach the wood above ground. Nests are built either in the soil underground or in wood<br />
above ground (Figure 15).<br />
In moist evergreen forest, 16 species of termite were as<br />
subterranean termites. One species belong to subfamily Rhinotermitinae, 3 species of<br />
subfamily Coptotermitinae, 7 species of subfamily Macrotermitinae, 3 species belong to<br />
subfamily Termitinae and 2 species of subfamily Apicotermitinae.<br />
Eleven species of termites in dry evergreen forest were<br />
subterranean termites. This group consists of 4 species belong to family<br />
Rhinotermitinae, 5 species of Macrotermitinae and 2 species belong to subfamily<br />
Termitinae.<br />
3.2.2 Arbore<strong>al</strong> nest<br />
56
Noirot (1970) described that this group usu<strong>al</strong>ly having nest which<br />
are constructed on trees, often at considerable heights, connected to the ground by<br />
covered g<strong>al</strong>leries which descend from the trunk of the tree where the nest is situated<br />
(Figure 16).<br />
Ten species of termite in moist evergreen forest were the termite<br />
lived in arbore<strong>al</strong> nest. This group consists of 1 species of Microcerotermes belong to<br />
Amitermitinae and 9 species belong to subfamily Nasutitermitinae.<br />
For dry evergreen forest, 11 species of termites were recorded as<br />
the termite lived in arbore<strong>al</strong> nest. This group consists of 1 species of Microcerotermes<br />
belong to Amitermitinae and 10 species belong to subfamily Nasutitermitinae.<br />
3.2.3 Epige<strong>al</strong> nest<br />
Bignell and Eggl<strong>et</strong>on (2000) explained that this group of termites<br />
usu<strong>al</strong>ly have colony centered on the ground of standing tree or against the side of trees.<br />
Their nests are usu<strong>al</strong>ly well defined and highly complex species specific structures but<br />
with a tendency to become more irregular as they age through erosion, additions and<br />
occupation by secondary inhabitants (Figure 16).<br />
In moist evergreen forest, 9 species of termite built epige<strong>al</strong> nests.<br />
This group consists of 2 species of Macrotermes (Macrotermitinae), 3 species belong to<br />
subfamily Amitermitinae, 1 species of Termes (Termitinae) and 3 species of<br />
Hospit<strong>al</strong>itermes belong to subfamily Nasutitermitinae.<br />
Five species of termite in dry evergreen forest built epige<strong>al</strong> nests.<br />
This group consists of 1 species of Macrotermes (Macrotermitinae), 3 species of<br />
Amitermitinae and 1 species of Termes, which belong to subfamily Termitinae.<br />
57
No. of species<br />
20<br />
15<br />
10<br />
5<br />
0<br />
4<br />
2<br />
<br />
<br />
10<br />
11<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
In wood Arbore<strong>al</strong> Epige<strong>al</strong> Subterranean<br />
Nest type<br />
MEF<br />
Similar to the reason explained earlier about subfamily K<strong>al</strong>otermitinae. Numbers<br />
of species of wood dwelling termites were found in higher degree in moist evergreen<br />
forest than in dry evergreen forest. Subfamily K<strong>al</strong>otermitinae classified into the wood<br />
dwellers that confined throughout of their life in wood consisted of dry-wood termite<br />
(Cryptotermes) and damp-wood termite (Gryptotermes) (Roonw<strong>al</strong>, 1970). The results<br />
which reve<strong>al</strong>ed that Glyptotermes was found in higher number of species in moist<br />
evergreen forest than dry evergreen forest might be due to the fact that in moist<br />
evergreen forest having higher average rainf<strong>al</strong>l and relative humidity than dry evergreen<br />
forest and may be more suitable for Glyptotermes, damp-wood termite, to live.<br />
Arbore<strong>al</strong> nesting termites were found in equ<strong>al</strong> number of species in moist<br />
evergreen forest and dry evergreen forest this might due to these forest types situated in<br />
region of heavy rainf<strong>al</strong>l. Lee and Wood (1971) reported that termite species that inhabit<br />
9<br />
5<br />
DEF<br />
Figure 13 Termite species in differing nest types in moist evergreen forest (MEF)<br />
and dry evergreen forest (DEF).<br />
16<br />
11<br />
58
egion of heavy rainf<strong>al</strong>l, especi<strong>al</strong>ly rain forest, maintenance of high humidity presents no<br />
problems, but protection from flooding and excessive erosion may be important. Many<br />
rain forest termites build arbore<strong>al</strong> nests, often with covered runways descending to the<br />
soil.<br />
Epige<strong>al</strong> nesting termites were found in higher number of species in moist<br />
evergreen forest than in dry evergreen forest. Three species of Hospit<strong>al</strong>itermes were<br />
epige<strong>al</strong> nesting termites which nested at the base of living tree and stump were found<br />
only in moist evergreen forest and Macrotermes were found in higher number of species<br />
in moist evergreen forest than in dry evergreen forest. Hesse (1955) found that the<br />
moisture content of inhabited mounds of three East African species of Macotermes was<br />
higher than that of adjacent uninhabited mound. The information above lead to the<br />
speculation that moist evergreen forest which having higher average rainf<strong>al</strong>l and relative<br />
humidity more than dry evergreen forest may be more suitable for epige<strong>al</strong> nesting<br />
termites to live.<br />
The results, which reve<strong>al</strong>ed that subterranean termite, were found in higher<br />
number of species in moist evergreen forest than in dry evergreen forest (Figure 13)<br />
might be related to report which showed that moist evergreen forest having more<br />
organic matter and soil moisture than dry evergreen forest (Sawasdee <strong>et</strong> <strong>al</strong>., 2001).<br />
Wachrinrat <strong>et</strong> <strong>al</strong>. (2001 a) reported that tot<strong>al</strong> aboveground biomass in moist evergreen<br />
forest higher than that of dry evergreen forest. The information above lead to the<br />
speculation that moist evergreen forest probably have more suitable soil organic matter,<br />
soil moisture and are b<strong>et</strong>ter feeding sites for subterranean termites to survive than dry<br />
evergreen forest.<br />
59
Figure 14 Wood-dwelling termites. A: Cryptotermes; B: Glyptotermes.<br />
A<br />
B<br />
60
Figure 15 Subterranean termites. A: Coptotermes; B: Ancistrotermes.<br />
A<br />
B<br />
61
A<br />
Figure 16 Arbore<strong>al</strong> nesting termites. A: Microcerotermes; B: Nasutitermes.<br />
62
Figure 17 Epige<strong>al</strong> nesting termites. A: Mound of Globitermes; B: Carton nest<br />
of Microcerotermes; C: Mound of Macrotermes; D: Nest of<br />
Termes.<br />
A<br />
D<br />
63
Diversity of Termite<br />
The results recorded from moist evergreen forest and dry evergreen forest were<br />
tabled and frequency of termites in cool-dry and hot-dry season of present study was<br />
further computed. Diversity of the ecosystem of termite in the study area were presented<br />
in Table 4 in term of frequency, number of species and <strong>al</strong>so H’ index.<br />
Table 4 Diversity of termites in moist evergreen forest and dry evergreen forest of<br />
Chanthaburi Province, Thailand during cool – dry and hot – dry season.<br />
Param<strong>et</strong>er<br />
Moist evergreen forest<br />
Cool-dry Hot-dry<br />
Dry evergreen forest<br />
Cool-dry Hot-dry<br />
Frequency (hits) of termites 243 222 231 234<br />
No. of species 31 21 24 17<br />
H’ index 2.415 2.284 2.121 1.67<br />
Result of diversity index (H’ index) as shown in Table 4 reve<strong>al</strong> that moist<br />
evergreen forest in cool-dry season showed the highest number of diversity index, which<br />
consistent with highest number of termite species found in moist evergreen forest.<br />
Dominant Species<br />
Table 5 showed the frequency of termites recorded in each type of forest<br />
ecosystem during cool-dry season and hot-dry season. Results from the study <strong>al</strong>so<br />
reve<strong>al</strong>ed the dominant species of termite in each forest type and season (Table 6 and<br />
Figure 18).<br />
64
Table 5 Frequency of termites recorded in each type of forest ecosystem during cooldry<br />
season and hot-dry season. MEF = moist evergreen forest, DEF = dry<br />
evergreen forest, F = number of family, SF = number of subfamily, G = number<br />
of genera, SP = number of species.<br />
Type of veg<strong>et</strong>ation<br />
Termite<br />
DEF MEF<br />
Genus and Species CoolHotCoolHotdrydrydrydry 1. F. K<strong>al</strong>otermitidae (2G, 4SP)<br />
1.1 SF. K<strong>al</strong>otermitinae 5 2 3 1<br />
1. Cryptotermes thailandis Ahmad 4 - 1 -<br />
2. Glyptotermes sp.1 - - 1 -<br />
3. Glyptotermes sp.2 1 2 - 1<br />
4. Glyptotermes sp.3 - - 1 -<br />
2. F Rhinotermitidae (2G, 6SP)<br />
2.1 SF Rhinotermitinae (1G, 1SP) 7 14 13 30<br />
5. Schedorhinotermes medioobscurus (Holmgren) 7 14 13 30<br />
2.2 SF Coptotermitinae (1G, 5SP) 4 3 4 8<br />
6. Coptotermes sp.1 - 2 - -<br />
7. Coptotermes gestroi Wasmann 2 - 3 2<br />
8. Coptotermes havilandi Holmgren - - - 1<br />
9. Coptotermes premrasmii Ahmad - - 1 5<br />
10. Coptotermes curvignathus Holmgren 2 1 - -<br />
3. F Termitidae (16G, 32SP)<br />
3.1 SF Macrotermitinae (5G, 9SP) 48 26 59 58<br />
11. Macrotermes annand<strong>al</strong>ei (Silvestri) 11 9 3 4<br />
12. Macrotermes maesodensis Ahmad - - 1 -<br />
13. Microtermes obesi Holmgren 3 - 1 2<br />
14. Ancistrotermes pakestanicus (Silvestri) 12 8 25 30<br />
15. Hypotermes makhamensis Ahmad 10 7 11 9<br />
16. Odontotermes sp.1 - - - 2<br />
17. Odontotermes proformosanus Ahmad 7 2 6 5<br />
18. Odontotermes formosanus (Shiraki) 5 - 11 6<br />
19. Odontotermes oblongathus Holmgren - - 1 -<br />
3.2 SF Amitermitinae (2G,4SP) 129 140 118 94<br />
20. Microcerotermes crassus Snyder 108 126 84 67<br />
21. Microcerotermes distans (Haviland) - 1 - -<br />
22. Microcerotermes annand<strong>al</strong>ei Silvestri - - 1 -<br />
23. Globitermes sulphureus (Haviland) 21 13 33 27<br />
3.3 SF Apicotermitinae (2G,2SP) - - 1 2<br />
24. Euhamitermes sp. - - - 1<br />
25. Speculitermes sp. - - 1 1<br />
3.4 SF Termitinae (4G, 4SP) 1 1 4 0<br />
26. Termes cosmis Haviland 1 - 2 -<br />
27. Pericapritermes sp.C - 1 - -<br />
28. Mirocapritermes concaveus Ahmad - - 1 -<br />
29. Angulitermes sp. - - 1 -<br />
65
Table 5 (cont.) Frequency of termites recorded in each type of forest ecosystem during<br />
cool-dry season and hot-dry season. MEF = moist evergreen forest,<br />
DEF = dry evergreen forest, F = number of family, SF = number of<br />
subfamily, G = number of genera, SP = number of species.<br />
Type of veg<strong>et</strong>ation<br />
Termite<br />
DEF MEF<br />
Genus and Species CoolHotCoolHotdrydrydrydry 3.5 SF Nasutitermitinae (3G, 13SP) 37 48 41 29<br />
30. Nasutitermes johoricus (John) 14 15 14 16<br />
31. Nasutitermes matangensiformis (Holmgren) 1 3 1 -<br />
32. Nasutitermes sp.1 3 1 1 -<br />
33. Nasutitermes sp.3 1 - - -<br />
34. Nasutitermes sp.4 1 - 1 -<br />
35. Nasutitermes sp.5 2 - 3 -<br />
36. Bulbitermes parapusillus Ahmad 9 17 9 3<br />
37. Bulbitermes laticeph<strong>al</strong>us Ahmad 2 12 4 5<br />
38. Bulbitermes sp.1 1 - 2 -<br />
39. Bulbitermes sp.2 3 - 5 -<br />
40. Hospit<strong>al</strong>itermes ataramensis Prashad and Sen-Sarma - - 1 -<br />
41. Hospit<strong>al</strong>itermes sp.3 - - - 4<br />
42. Hospit<strong>al</strong>itermes sp.4 - - - 1<br />
Table 6 Frequency of dominant species in moist evergreen forest and dry evergreen<br />
forest during cool-dry season and hot-dry season.<br />
Dominant species<br />
MEF<br />
Cool-dry Hot-dry<br />
DEF<br />
Cool-dry Hot-dry<br />
1. Microcerotermes crassus Snyder 84 67 108 126<br />
2. Globitermes sulphureus (Haviland) 33 27 21 13<br />
3. Ancistrotermes pakestanicus (Silvestri) 25 30 12 8<br />
4. Schedorhinotermes medioobscurus<br />
(Holmgren)<br />
13 30 7 14<br />
66
Frequency of termite<br />
140<br />
120<br />
100<br />
80<br />
60<br />
40<br />
20<br />
0<br />
126<br />
108<br />
<br />
<br />
84<br />
<br />
<br />
67<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
33 27 25 30 30<br />
<br />
<br />
21<br />
<br />
13 <br />
12 <br />
<br />
<br />
<br />
8 13<br />
7<br />
14<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
Microcerotermes crassus Globitermes sulphureus Ancistrotermes<br />
pakestanicus<br />
Dominant species<br />
M cool-dry<br />
M hot-dry<br />
D cool-dry<br />
D hot-dry<br />
<br />
<br />
Schedorhinotermes<br />
medioobscurus<br />
Figure 18 Frequency of dominant species in moist evergreen forest and dry<br />
evergreen forest during cool-dry season and hot-dry season.<br />
In this study, the dominant species was d<strong>et</strong>ermined from the frequency of<br />
termites found within the study areas. Any species found over 20 spots from each area<br />
were considered as dominant species. Result obtained from the mentioned<br />
characteristic reve<strong>al</strong>ed that Microcerotermes crassus Snyder (Figure 19 A) is dominant<br />
species in both moist evergreen forest and dry evergreen forest. Moist evergreen forest<br />
in cool-dry season showed the frequency of this species more than hot-dry season. On<br />
the contrary, dry evergreen forest in cool-dry season showed the frequency less than<br />
hot-dry season.<br />
The results <strong>al</strong>so reve<strong>al</strong>ed that Microcerotermes crassus Snyder was found as a<br />
dominant species in both of moist evergreen forest and dry evergreen forest. This might<br />
be due to the fact that this termite has adaptive ability to sever<strong>al</strong> conditions.<br />
67
Globitermes sulphureus (Haviland) (Figure 19 B) is a secondary dominant<br />
species in moist evergreen forest both of 2 seasons but in dry evergreen forest this<br />
species is a secondary dominance only in cool-dry season.<br />
According to the results shown in Figure 18 reve<strong>al</strong>ed that Globitermes<br />
sulphureus (Haviland) is a secondary dominant species in moist evergreen forest both<br />
of 2 seasons. Ahmad (1965) mentioned that this termite is one of the most common<br />
mound-building termites of Thailand.<br />
Ancistrotermes pakestanicus Silvestri (Figure 19 C), the fungus growing<br />
termites, were found to be another secondary dominant species particularly in moist<br />
evergreen forest, they were <strong>al</strong>so found in dry evergreen forest in both seasons but in<br />
less frequency which was not enough to be considered as dominance.<br />
Schedorhinotermes medioobscurus (Holmgren) (Figure 19 D) were found as<br />
dominance only in hot-dry season of moist evergreen forest (30 times were found).<br />
68
0.5 mm. A 0.5 mm.<br />
B<br />
0.5 mm.<br />
C<br />
1.0 mm.<br />
Figure 19 Dominant species in moist evergreen forest and dry evergreen forest.<br />
A: Soldier of Microcerotermes crassus (dominant species in 2 forest<br />
types); B: Soldier of Globitermes sulphureus (secondary dominant in<br />
moist evergreen forest); C: Soldier of Ancistrotermes pakestanicus<br />
(secondary dominant in moist evergreen forest); D: Soldier of<br />
Schedorhinotermes medioobscurus.<br />
D<br />
69
Termite Species in Relation to Season<strong>al</strong> Changes<br />
Results of survey as shown in Table 7 and Figure 20 reve<strong>al</strong>ed that in moist<br />
evergreen forest, the number of species in subfamily K<strong>al</strong>otermitinae, Amitermitinae,<br />
Macrotermitinae, Termitinae and Nasutitermitinae that tend to decrease in hot-dry<br />
season. While the number of species in subfamily Coptotermitinae and Apicotermitinae<br />
increase in hot-dry season. There is no difference in the number of species in subfamily<br />
Rhinotermitinae in 2 seasons.<br />
In dry evergreen forest, the number of species in subfamily K<strong>al</strong>otermitinae,<br />
Macrotermitinae and Nasutitermitinae tend to decrease in hot-dry season while the<br />
number of species in Amitermitinae increase in hot-dry season. There is no difference in<br />
the number of species in subfamily Rhinotermitinae, Coptotermitinae and Termitinae in 2<br />
seasons.<br />
Table 7 Number of species of termites recorded in each type of forest ecosystem<br />
during cool-dry season and hot-dry season.<br />
Number of species<br />
Subfamily<br />
Moist evergreen forest Dry evergreen forest<br />
Cool-dry Hot-dry Cool-dry Hot-dry<br />
K<strong>al</strong>otermitinae 3 1 2 1<br />
Rhinotermitinae 1 1 1 1<br />
Coptotermitinae 2 3 2 2<br />
Macrotermitinae 8 7 6 4<br />
Amitermitinae 3 2 2 3<br />
Apicotermitinae 1 2 0 0<br />
Termitinae 3 0 1 1<br />
Nasutitermitinae 10 5 10 5<br />
70
No. of species<br />
12<br />
10<br />
8<br />
6<br />
4<br />
2<br />
0<br />
K<strong>al</strong>otermitinae<br />
3<br />
2<br />
1 1 1 1 1 1<br />
<br />
<br />
<br />
<br />
<br />
<br />
Rhinotermitinae<br />
Coptotermitinae<br />
<br />
M cool-dry<br />
3 3 3<br />
2 2 2 2 2<br />
<br />
<br />
<br />
<br />
<br />
Amitermitinae<br />
<br />
<br />
<br />
<br />
<br />
Macrotermitinae<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
8<br />
7<br />
6<br />
4<br />
3<br />
1 1<br />
0<br />
2<br />
1<br />
0 0<br />
Termitinae<br />
Subfamily<br />
<br />
<br />
M hot-dry D cool-dry<br />
Apicotermitinae<br />
<br />
D hot-dry<br />
Results indicated that species diversity and species richness of termite in moist<br />
evergreen forest and dry evergreen forest decreased in hot-dry season. Soil moisture<br />
has much more effect to species richness and termite activity than temperature and<br />
relative humidity (Pearce, 1999). In eastern Thailand the climatic condition particularly<br />
temperature and relative humidity among season is not much different excepted for<br />
rainf<strong>al</strong>l (Wachrinrat <strong>et</strong> <strong>al</strong>., 2001 a). Report on volume of water content at depth 0-15 cm<br />
in moist evergreen forest and dry evergreen forest of the study site were 0.207 and<br />
0.173 respectively (Sawasdee <strong>et</strong> <strong>al</strong>., 2001). During the survey in hot-dry season (April)<br />
there was heavy rainf<strong>al</strong>l and might have decreased termite richness and activity in this<br />
study.<br />
10<br />
<br />
<br />
Nasutitermitinae<br />
Figure 20 Number of species of termites recorded in each type of forest<br />
ecosystem during cool-dry season and hot-dry season.<br />
10<br />
<br />
<br />
<br />
<br />
5 5<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
<br />
71
CONCLUSION<br />
1. Three hundred and forty five samples of termite were collected from moist<br />
evergreen forest at Khao Kitchagoot Nation<strong>al</strong> Park and dry evergreen forest at Khao Soi<br />
Dao Wildlife Sanctuary identification of surveyed samples resulted three families, 8<br />
subfamilies, 20 genera and 42 species.<br />
2. Results of the survey reve<strong>al</strong>ed that moist evergreen forest showed the higher<br />
number of termite species than dry evergreen forest (37 species of 19 genera to 27<br />
species of 15 genera).<br />
3. Moist evergreen forest having higher average rainf<strong>al</strong>l and relative humidity<br />
than dry evergreen forest and might be suitable for termite in subfamily K<strong>al</strong>otermitidae,<br />
Macrotermitidae and Nasutitemitinae to establish their colony.<br />
4. Moist evergreen forest probably has more appropriate soil organic matter<br />
and soil moisture for termites in subfamily Apicotermitinae and Termitinae to survive than<br />
dry evergreen forest.<br />
5. There is no different in the number of species of subfamily Rhinotermitinae,<br />
Coptotermitinae and Amitermitinae in moist evergreen forest and dry evergreen forest. It<br />
was therefore, assumed that these two forest types probably contain sufficient amount of<br />
trees, wood and litterf<strong>al</strong>l for these termites to forage.<br />
6. Classification of termites base on their food habitat resulted into four different<br />
termite group as followed:<br />
6.1 Wood feeders<br />
6.2 Wood and leaf feeders<br />
6.3 Soil or humus feeders<br />
72
6.4 Lichen feeders<br />
7. Classification of termites base on their nest habitat resulted into two major<br />
groups as followed:<br />
7.1 Wood dwellers<br />
7.2 Ground dwellers; The ground dwellers can be further classified into 3<br />
sub-groups:<br />
7.2.1 Subterranean termites<br />
7.2.2 Arbore<strong>al</strong> nesting termites<br />
7.2.3 Epige<strong>al</strong> nesting termites<br />
8. Number of diversity index (H’ index) of moist evergreen forest was higher<br />
than dry evergreen forest in both cool-dry and hot-dry seasons. And, consequently<br />
number of diversity index of moist evergreen forest was found to be highest in cool-dry<br />
season.<br />
9. Microcerotermes crassus Snyder was found as first dominant species in<br />
both moist evergreen forest and dry evergreen forest. Globitermes sulphureus<br />
(Haviland) is the secondary dominant species in both moist evergreen forest and dry<br />
evergreen forest. Ancistrotermes pakestanicus (Silvestri) which is a fungus growing<br />
termites reve<strong>al</strong>ed themselves as another dominant species in moist evergreen forest.<br />
10. Identification of specimen from different time of survey reve<strong>al</strong>ed that<br />
season<strong>al</strong> changed did not have much effect to species richness of termite in <strong>al</strong>l study<br />
sites. Species diversity and species richness or termite activity tend to be lower in hotdry<br />
season (April).<br />
73
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APPENDIX<br />
83
Appendix Table 1 The data recording form used in the field work to d<strong>et</strong>ermine ecologic<strong>al</strong> niches of termites.<br />
Date<br />
Plot No<br />
Termite<br />
species<br />
Nest on<br />
living<br />
tree<br />
Nest on<br />
ground<br />
Mound<br />
Tunnel<br />
on living<br />
tree<br />
Tree<br />
stump<br />
Wood<br />
debris<br />
Leave<br />
litter<br />
log In soil<br />
Mushroom<br />
fungus<br />
garden<br />
Sign of<br />
termite<br />
attack<br />
Enemies<br />
Remark<br />
Termite Present<br />
many little<br />
Case
Appendix Table 2 Climatic data in moist evergreen forest and dry evergreen forest of<br />
Chanthaburi Province 1/ .<br />
Climatic data Moist evergreen forest Dry evergreen forest<br />
Average annu<strong>al</strong> rainf<strong>al</strong>l (mm.) 2,865 mm. 1,517 mm.<br />
Relative humidity (%) 79% 73%<br />
Average temperature (°C) 27 °C 27 °C<br />
Data from : 1/ Wachrinrat <strong>et</strong> <strong>al</strong>. (2001 a)<br />
Appendix Table 3 The volume of water content, percentage of organic matter and soil<br />
litter surface thickness in moist evergreen forest and dry evergreen<br />
forest 1/ .<br />
Param<strong>et</strong>er Moist evergreen forest Dry evergreen forest<br />
Volume of water content<br />
(depth 0-15 cm)<br />
0.207 0.173<br />
Organic matter<br />
(depth 0-15 cm)<br />
5.55 3.99<br />
Soil litter thickness (cm.) 7.40 5.68<br />
Data from : 1/ Sawasdee <strong>et</strong> <strong>al</strong>. (2001)<br />
85
Appendix Table 4 Dominant trees in moist evergreen forest and dry evergreen forest 1/ .<br />
Dominant Trees Moist evergreen forest Dry evergreen forest<br />
- F. ANNONACEAE<br />
1. Alphonsea elliptica Hook.f.&Thoms. - ✓<br />
- F. CEASALPINIACEAE<br />
2. Saraca declinata Miq. - ✓<br />
- F. EBENACEAE<br />
3. Diospyros dictyoneura Hiern. - ✓<br />
4. Diopyros pendula Hasselt ex Hassk. ✓ -<br />
5. Diospyros rubra Lec. - ✓<br />
- F. ICACINACEAE<br />
6. Gonocaryum lobbianum Kurz<br />
- F. LAURACEAE<br />
7. Crytocarya scortechnii - ✓<br />
8. Cyathoc<strong>al</strong>yx martabanicus Hook.f.&th. - ✓<br />
- F. MELIACEAE<br />
9. W<strong>al</strong>sura robusta Roxb. - ✓<br />
- F. MIMOSACEAE<br />
10. Archidendron guocense (Pierre) ✓ -<br />
- F. MYRISTICACEAE<br />
11. Knema globularia Warb. ✓ -<br />
12. Knema laurina Warb. ✓ -<br />
- F. MYRTACEAE<br />
13. Cleistoc<strong>al</strong>yx operculatus Merr.& Perry ✓ -<br />
14. Syzygium cf. diospyrifolium S.N.Mitra ✓ -<br />
15. Syzygium lineatum Merr.& L.M.Perry ✓ -<br />
- F. OLACACEAE<br />
16. Strombosia javanica Bl. ✓ ✓<br />
- F. SAPINDACEAE<br />
17. Xerospermum laevigatum Radlk. ssp. - ✓<br />
- F. STERCULIACEAE<br />
18. Pterygota <strong>al</strong>ata R.Br. - ✓<br />
19. Scaphium macropodum Beaumee ✓ -<br />
Data from : 1/ Wachrinrat <strong>et</strong> <strong>al</strong>. (2001 a)<br />
86
Appendix Table 5 Tot<strong>al</strong> aboveground biomass and tot<strong>al</strong> litterf<strong>al</strong>l in moist evergreen<br />
forest and dry evergreen forest, Chanthaburi Province.<br />
Param<strong>et</strong>er Moist evergreen forest Dry evergreen forest<br />
Tot<strong>al</strong> Biomass (ton. ha -1 ) 1/<br />
434.732 340.558<br />
Tot<strong>al</strong> litterf<strong>al</strong>l (kg. ha -1 ) 2/<br />
6,627.85 7,771.63<br />
Data from : 1/ Wachrinrat <strong>et</strong> <strong>al</strong>. (2001 a)<br />
2/ Wachrinrat <strong>et</strong> <strong>al</strong>. (2001 b)<br />
Appendix Table 6 Quatitative characteristics of each sample plot in evergreen forest at<br />
Chanthaburi Province, only trees with DBH ≥ 4.5 m. are included 1/ .<br />
Param<strong>et</strong>er Moist evergreen forest Dry evergreen forest<br />
No. of species (sp.) 135 138<br />
No. of individu<strong>al</strong>s (no.ha -1 ) 1,510 1,355<br />
Mean DBH (cm) 13.67 13.07<br />
Mean height (m) 11.45 9.54<br />
Bas<strong>al</strong> area (%) 0.4799 0.3995<br />
Data from : 1/ Wachrinrat <strong>et</strong> <strong>al</strong>. (2001 a)<br />
87