Species Richness, Functional Diversity And Relative Abundance Of Termites Under Different Land Use Regimes
“Best bet” Land-use Systems
Thematic reports
Impact of different land uses on biodiversity
An Intensive Biodiversity Baseline Study in Jambi Province,Central Sumatra, Indonesia
Unique id: 8
Source file: D:\Projects\ASB\ASB Country and Thematic reports - xml\Above ground biodiversity assessmet WG\C-Sec8-9.xml
Authors: D.T. Jones, F.X. Susilo, D.E. Bignell, H. Suryo
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8.1 Introduction:
Much of
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).
8.2 Aims:
To
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.
The search
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
8.3 Personnel:
Principal Investigator:
Dr D.T. Jones (termite ecologist and
taxonomist) - Biodiversity Division Entomology Department
The
Assisted by:
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,
8.4 Methods:
Seven sites in
Land use type Site code Date sampled
1. Paraserianthes
plantation BS 6 19
+ 20 Nov. 1997
2. Primary forest BS 1 21 + 22 Nov. 1997
3. Logged-over forest BS
3 22 + 23 Nov. 1997
4. Imperata
grassland BS 12
24 Nov.
1997
5. Cassava garden BS 14 24 + 25 Nov. 1997
6. Jungle rubber BS 10 26 + 27 Nov. 1997
7. Rubber plantation BS
8 27 + 28 Nov. 1997
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)
Functional groups:
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.
8.6 Discussion:
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
Conclusions:
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
Acknowledgements
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
while in
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Table 8.1Species
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
Primary Logged Jungle Rubber Parase-Imperata Cassava
Species Functional forest forest rubber pltn. ianthes
grassland garden
group (BS 1) (BS 3) (BS 10) (BS 8) (BS 6) (BS 12)
(BS 14)
KALOTERMITIDAE
Glyptotermes sp. W - - - 1 -
- -
RHINOTERMITIDAE
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 -
- -
Primary Logged Jungle Rubber Parase-Imperata Cassava
Species Functional forest forest rubber pltn. ianthes
grassland garden
group (BS 1) (BS 3) (BS 10) (BS 8) (BS 6) (BS 12)
(BS 14)
TERMITIDAE
Macrotermitinae
Macrotermes gilvus W - - - - - - 1
Macrotermes sp. 1 W 1 - - - - - -
Odontotermes denticulatus W - - 5 - - - -
Odontotermes sarawakensis W 10 9 - - - - -
Ancistrotermes pakistanicus W - - 3 - - - -
Termitinae
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 - - - - -