Species Richness, Functional Diversity And Relative Abundance Of Termites Under Different Land Use Regimes
“Best bet” Land-use Systems
Impact of different land uses on biodiversity
An Intensive Biodiversity Baseline Study in Jambi Province,Central Sumatra, Indonesia
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Authors: D.T. Jones, F.X. Susilo, D.E. Bignell, H. Suryo
Within the context of sustainable
agricultural production under conditions of rapid land-use change, declining
forest cover, loss of biodiversity and an increasing human population, research
should be focused on those groups of organisms that contribute directly to
plant productivity and their response to changes in land use. The importance of
invertebrate macrofauna to the promotion of tropical soil fertility has been
stressed in recent reviews (Fragoso et
al., 1993; Lavelle et al., 1994;
Garnier-Sillam & Harry, 1995; Nash & Whitford, 1995; Brussaard &
Jumas, 1996; Wood, 1996). The
distribution, protection and stabilization of organic matter, the genesis of
soil structure, humification, the release of immobilized N and P, the
improvement of drainage and aeration, and the increase in exchangeable cations
have all been demonstrated in soils modified by termites and earthworms
(Lavelle et al., 1997). In African systems, forest clearance depletes
termite abundance and diversity (Wood et
al. 1982; Eggleton et al.,1995;
1996) but similar studies are not yet available from
Termites are a key functional group
of animals in the tropics and can achieve very high populations. For example, in the forests of southern
Given the ecological importance of termites, there is a need to characterize termite assemblage structure within and between sites. As a consequence of their highly patchy spatial distribution, combined with the many and varied field sampling regimes adopted by previous researchers, it has not been possible to use the existing data to make reliable direct comparisons of termite diversity and abundance between sites (see Eggleton & Bignell, 1995). As Sutton & Collins (1991) emphasised, it is necessary to develop and test standardised sampling methods that can be applied easily throughout the tropics. To this end, a standardised transect sampling method designed to measure termite species richness and functional diversity in tropical forests has been developed. The protocol has been used in Cameroon (Eggleton et al, 1995), Thailand (Davies, 1997), Peninsular Malaysia (Jones & Brendell, in press) and two sites in Sabah; Maliau Basin (Jones et al., in press) and Danum Valley (Eggleton et al., in press).
assess the termite assemblage under different land uses. The first aim is to
measure species richness, functional diversity and the relative abundance of
termites under seven different land-use regimes in
The responses of termites to land-use changes. If the history of exploitation at each of the Jambi study sites is known, it may be possible to arrange the sites along a 'land-use intensification gradient' or into one or more 'land-use sequences'. By assuming that all the Jambi sites were originally forested and had similar termite assemblages, it will be possible to hypothesise about the response of termites to changes in land use. This assumes the primary assumption is correct.
for correlates between termites and other organisms. The multidisciplinary
approach adopted in this project is rare in ecological field studies. In all,
seven groups of organisms have been studied in the same sites in
Dr D.T. Jones (termite ecologist and taxonomist) - Biodiversity Division Entomology Department
Dr David Bignell (termite physiologist), Tropical Biology & Conservation Unit, University Malaysia Sabah.
Dr F.X. Susilo (entomologist) - Jurusan Proteksi Tanaman, Fakultas Pertanian, Universitas Lampung
Dr Suryo Hardiwibowo (biologist) -
Gadjah Mada Universitas,
Seven sites in
The standardized transect sampling method:
All transects were co-located with a 40x5m strip transect used to sample vegetation and for other multidisciplinary studies. Each termite transect was 100 m long and 2 m wide, divided into 20 contiguous sections (each 5 m x 2 m), and numbered sequentially. Each section was sampled by two people for 30 minutes (a total of one hour of collecting per section). In order to standardise sampling effort, the trained collectors worked steadily and continuously during the 30 minute collecting period. In each section the collectors searched the following microhabitats which are common sites for termites: surface soil to 5 cm depth; accumulations of litter and humus at the base of trees; the inside of dead logs, tree stumps, branches and twigs; the soil within and beneath very rotten logs; all subterranean nests, mounds, carton sheeting and runways on vegetation, and arboreal nests up to a height of 2 m above-ground level.
The protocol was designed to offer a flexible approach to the sampling, whereby the collectors used their experience and judgement to search for, locate and sample as many species of termite in each section as time allowed. Specimens from each termite population encountered were sampled. All castes were collected if present, but priority was given to finding soldiers and workers. Termites were placed in vials labelled with the section number and filled with 80%ethanol.
In structurally complex habitats, i.e. with a relatively large above-ground biomass (such as forests and plantation systems), the collectors spend approximately half their collecting time searching the above-ground microhabitats described above. The remaining 15 minutes were used searching for termites in the soil. However, in the case of the Imperata grassland (BS 12) and the Cassava garden (BS 14) there was relatively little above-ground biomass. Within the transects in both systems there were no trees and virtually no dead wood or leaf litter. Therefore, in these land-use types (Imperata grassland and the Cassava garden) the collectors sampled only for 15 minutes (total collecting effort = 30 minutes) in each section. This procedure ensured that equal effort was given to searching for termites in the soil in each transect.
The transect sampling method provides a semi-quantitative measure of the relative abundance of termites based on the number of encounters or 'hits' with each species in a transect. A hit is defined as the recorded presence of a species in one section. Therefore, if a species is present in every section of a transect it will have a relative abundance score of 20. The number of hits per transect can then be used as an indicator of the relative abundance of termites occurring within a transect, as well as between transects. It gives no measure of the absolute abundance per unit area.
8.4.2 Identification of material:
During the field trip, great effort was taken to examine as much of the material as time allowed. This was made possible due to the microscope and light source provided by David Bignell. In the evenings, many hours were spent making provisional identifications. All samples with soldiers were identified to genus, and then morphospecies numbers were allocated. A working reference collection was maintained so that material from all transects could be cross-referenced and the morphospecies designations applied consistently. Many vials contained two or more species, and some of these were separated where time and accuracy allowed. Two groups of samples were not identified. The first were samples with workers (i.e. no soldier specimens collected). Workers are difficult and time consuming to identify as the mandibles must be dissected, and the structure of the gut must be examined, sometimes necessitating the removal and mounting of the enteric valve. The second group were genera in the Subulitermes complex. These are small termites whose taxonomy is ill-defined and that are difficult to identify.
It must be stressed that the results given in this report are based solely on the provisional identifications made during the field trip. At the Natural History Museum every sample will be examined again, and accurate species-level identifications will be made. By comparison with the museum's extensive reference collection (which contains approximately 16000 vials of identified material, plus about 1000 vials of type material), it will be possible to put specific names on alarge proportion of the Jambi collection. It is estimated that the identification work at the museum should take about 4 to 5 weeks [note: completed July 1998. eds. See Annex III, Table 11)
Genera were assigned to one of five functional groups based on known feeding habits (see Collins, 1984; Eggleton et al., 1996; Jones et al., in press; Eggleton et al., submitted), the shape of the molar plates of the worker mandibles (Deligne, 1966), and worker gut content analyses (Sleaford et al., 1996). The functional groups are;
Soil-feeding: termites that feed on humus and mineral soil.
Wood-feeding: termites that feed on dead wood.
Soil/wood interface-feeding: termites that feed on extremely decayed wood that has lost its structure and become soil-like.
Litter-feeding: termites that feed exclusively on leaf-litter and small items of woody trash.
Epiphyte-feeding: Hospitalitermes is known to feed on lichens and other free living non-vascular plants which they graze from the surface of tree trunks (Collins, 1979; Jones & Gathorne-Hardy, 1995).
8.5 Preliminary results:
8.5.1 Species richness:
The preliminary sorting carried out
during the Jambi field work produced a conservative total of 23 genera and 48
morphospecies for all seven land-use types (Annex III, Table 11). However, in addition to these taxa, the
Subulitermes complex and many vials of workers await examination. The senior author speculates that these vials
will possibly contain several genera plus between 3 to 10 species which can be
added to the checklist. Members of the
Apicotermitinae subfamily are rare inSoutheast Asia but have been collected in
transects run in
Table 8.1 gives the list of morphospecies currently recorded from each transect. The preliminary identifications clearly show that the primary forest site is the most species rich, while the Imperata grassland and the Cassava garden sites are the most depauperate. The logged-over forest site and the agroforestry systems all have intermediate levels of species richness. Figure 8.1 displays the taxonomic composition of each transect sample. The Termitinae are the dominant subfamily in sites except the Paraserianthes plantation site and the Cassava garden system.
8.5.2 Relative abundance:
The number of hits (the presence of a species in a section) is recorded in Table 8.1. Termites are most abundant in the primary forest site and least abundant in the Cassava garden. The termites collected in this study fall into four feeding groups. Wood-feeding and soil-feeding species are relatively abundant in most transects, while epiphyte-feeders are rare and interface-feeders (those species that feed on extremely decayed soil-like wood) vary considerably in abundance among transects. Figure 8.2 displays the relative abundance of termites in each functional group. Of notable interest is the high relative abundance of soil-feeders in the jungle rubber system, and their absence from the Paraserianthes plantation. Grass-harvesting species and taxa that feed exclusively on leaf-litter appear to be absent from the study sites.
It must be stressed that the results given in the table and figures are based on provisional identifications. Table 8.1 also lists the number of vials containing specimens of the Subulitermes complex and workers which still await examination, and suggest the possible extent of extra species and hits that may be added to each transect. While we are certain that the final results for most of the transects will vary in species richness and relative abundance from those presented here, the senior author is confident that the overall patterns are likely to be similar to those already evident in the preliminary results.
Our knowledge of the termite fauna of
The transect method has been tested
against known local termite faunas and shown to produce representative samples
that are not significantly different in taxonomic or functional composition
from their local assemblage (Jones & Eggleton, in prep.). The highest
species richness found in Southeast Asian forests using the transect method is
33 species at
The preliminary results show a decline in termites species richness (Fig. 8.1) and relative abundance (Fig. 8.2) across the seven land-use types. Casual observations of the botanical features at each site by the authors suggested a positive relationship between termite species richness and physical complexity. It has been speculated that the degree of canopy closure appears to have a strong influence on termite diversity (Eggleton et al., 1995, 1996). Preliminary results from Jambi show a very high correlation between termite relative abundance and the recorded basal area of woody plants (r2 = 0.95; Gillison, pers. comm.; see also Annex II, Figure 1c). We await the dissemination of the vascular plant data to investigate whether there are significant correlates between the termite assemblages and the plant communities.
The efficiency of the transect
method, based on the number of species collected per unit effort (number of
days for one trained person to collect and identify samples) has already been
calculated (Jones & Eggleton, in prep.). One transect takes one trained
collector four days to complete. The
material from one primary forest transect at
With the completion of seven termite
transects and the preliminary sorting, the field-based phase of the Jambi
project can be considered a great success.
When all the museum-based identification work is complete, the top set
of material will be deposited at the
The authors would like to thank Dr
Andy Gillison and Ir Nining Liswanti for organising the field work in Jambi and
all the travel arrangements. In
addition, we are grateful to the logistical support provided by CIFOR and ICRAF
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checklist of termites collected from the seven land-use types in
Termites were collected using the standardised transect sampling protocol. One transect was run in each land-use type. Figures are the relative abundance of each species, based on the number of 'hits' of each species in a transect (the presence of a species in one section represents one hit). Functional group are: W = wood-feeders, I = soil/wood interface-feeders, S = soil-feeders, E = epiphyte-feeders
Species Functional forest forest rubber pltn. ianthes grassland garden
Glyptotermes sp. W - - - 1 - - -
Coptotermes curvignathus W 1 1 1 3 1 - -
Coptotermes sepangensis W - - - - 4 - -
Coptotermes borneensis W - - - - 1 - -
Heterotermes tenuior W 1 - - - - - -
Parrhinotermes near minor W - - 1 - - - -
Parrhinotermes near sp. C W - 1 - - - - -
Schedorhinotermes javanicus W 1 - 7 - 7 - -
Schedorhinotermes sarawakensis W 1 - - - 9 - -
Schedorhinotermes tarakanensis W 6 7 4 1 - - -
Schedorhinotermes sp. W - - - 2 - - -
Species Functional forest forest rubber pltn. ianthes grassland garden
Macrotermes gilvus W - - - - - - 1
Macrotermes sp. 1 W 1 - - - - - -
Odontotermes denticulatus W - - 5 - - - -
Odontotermes sarawakensis W 10 9 - - - - -
Ancistrotermes pakistanicus W - - 3 - - - -
Prohamitermes mirabilis I 3 7 - 6 4 - -
Labritermes buttelreepeni S - - 1 2 - - -
Globitermes globosus W 8 4 1 - - 4 -
Microcerotermes serrula W 3 7 - 1 - - -
Microcerotermes near havilandi W - 1 - - - - -
Termes comis I 4 1 - - 1 - -
Termes propinquus I 3 - - 12 1 -
Homallotermes eleanorae I 1 - - 3 - - -
Homallotermes foraminifer I 1 4 - - - - -
Mirocapritermes connectens S - 2 10 - - - -
Malaysiocapritermes prosetiger S 3 2 10 - - - -
Table 8.1Species checklist of termites
collected from the seven land-use types in
Species Functional forest forest rubber pltn. ianthes grassland garden
group (BS 1) (BS 3) (BS 10) (BS 8) (BS 6) (BS 12) (BS 14)
Procapritermes sandakanensis S - - 3 - - - -
Procapritermes setiger S 8 6 2 - - - -
Procapritermes near minutus S 4 - 1 - - - -
Procapritermes sp. A S - - 5 - - - -
Coxocapritermes sp. A S 6 1 - - - - -
Coxocapritermes sp. C S 2 3 - - - - -
Coxocapritermes sp. D S 1 3 2 - - - -
Kemneritermes sp. A S 4 1 - - - - -
Pericapritermes dolichocephalus S - - 6 - - - -
Pericapritermes nitobei S 1 - 2 - - - -
Pericapritermes semarangi S 2 - - - - 5 -
Dicuspiditermes nemorosus S 11 18 12 12 - - -
Dicuspiditermessantschii S 6 5 1 2 2 - -
Havilanditermes proatripennis W - - - 6 - - -
Nasutitermes havilandi W 1 - 2 - 3 - -
Nasutitermes matangensiformis W - - 2 - - - -
Nasutitermes neoparvus W - - - 1 - - -
Nasutitermes sp. C W - - - 2 - - -
Nasutitermes sp. D W 1 - - - 2 - -
Species Functional forest forest rubber pltn. ianthes grassland garden
Bulbitermes germanus W 2 - - - - - -
Bulbitermes prabhae W 1 - - - - - -
Bulbitermes sp. A W 3 1 - - - - -
Hospitalitermes hospitalis E 4 - - 2 - - -
Hospitalitermes sp. G E - - - - - - -
Proaciculitermes ?malayanus S 1 3 - - - - -
Proaciculitermes sp. B S 2 3 - - - - -
Aciculioiditermes sp. C S 1 - - - - - -
Oriensubulitermes inanis S 2 4 2 - - - -
Number of species 35 23 22 16 11 2 1
Figure 8.1. Species richness of termites collected from transects in seven land-use types
Figure 8.2. Relative abundance of termites collected from transects in seven land-use