Article history Received:. Cite Icon Cite. Abstract A comparative study of termite Isoptera: Termitidae species diversity and richness in three ecologically different habitats was conducted in the Shivamogga District of Karnataka, India, from September to August You do not currently have access to this content.
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Burkness, William D. Hutchison, Matthew D. Randell, Phillip M. Roberts, A. Stanley Culpepper. Logged forest and oil palm plantations now dominate the landscape of Malaysian Borneo Bryan et al. Although selectively logged forests retain many species e. Berry et al. For example, a review of bird responses to tropical forest disturbance Gray et al.
Also, a review of tropical forest dung beetle communities showed similar diversity declines with increasing habitat disturbance, along with a reduction in the number of forest species Nichols et al. A range of taxa including birds Peh et al.
Changes in assemblages, and particularly the loss of functionally important species, can have significant impacts on ecosystem functioning Hooper et al. Termites and ants are among the most important insect groups in tropical forest ecosystems.
Termites feed on plant material in varying stages of decay e. They play major roles in processes such as decomposition, and nutrient and carbon cycling Eggleton et al.
Ants disperse seeds, assist soil processing and nutrient cycling, and are mutualists with a range of species e. As both of these social insect groups play substantial ecological roles, the potential for interaction between them is important. Many ants feed on termites, and some ant species are specialised termite feeders e.
Mutualistic interactions between ants and termites, such as nest-sharing, have also been observed Jaffe et al. In addition to direct predatory and mutualistic interactions, ants and termites may interact indirectly through changes they make to their environments. Both groups are major ecosystem engineers Jones et al. Folgarait ; Lavelle and Spain ; Jouquet et al. Given the major roles of ants and termites in ecosystem function it is likely that functioning and resilience of both rain forest and oil palm plantation ecosystems will be affected by the abundance and composition of ant and termite assemblages Naeem et al.
Previous studies have shown that both ant and termite diversity usually decrease following habitat conversion Jones et al. Termites and ants also show shifts in assemblage structure with habitat disturbance. Soil feeding termites are vulnerable to loss of old growth forest, although wood feeders may have more species in mature regenerating forest Eggleton et al.
We know of no studies that have either, a sampled ants and termites simultaneously across a forest disturbance gradient or, b considered termite community composition in oil palm plantation. Here we assess the co-variation in functional and feeding group composition of ants and termites along a habitat disturbance gradient comprising sites in old growth forest, logged forest and oil palm plantation converted from logged forest, in Sabah, Malaysian Borneo.
All sampling was conducted in Sabah, Malaysian Borneo, at an average of m asl. Old growth forest survey points at Maliau were in forest that has never been logged commercially, although half of the survey points were in forest that has been lightly logged once. Stand basal area in this lightly-logged area remains similar to undisturbed sites Hamzah Tangki, unpublished data and substantially different from the commercially logged forest Ewers et al.
Tree communities were deemed not to have changed significantly Ewers et al. Logged forest survey points were in forest that has been selectively logged twice: once during the s and again from the late ss. Oil palm plantation survey points were in areas of Elaeis guineensis monocultures, planted in 10 years old at time of survey , with a low, open canopy and sparse understory vegetation.
Further details are provided in Ewers et al. Fifty-nine survey points were sampled in our study: 18 in old growth forest, 32 in logged forest of varying forest quality, and nine in oil palm plantation Online Resource, Fig.
A larger number of survey points were sampled in logged forest and old growth forest because we expected these habitats to be more heterogeneous and we wanted our points to span a gradient of habitat disturbance across all the habitats. Neighbouring survey points were m apart. Selection of these survey points was made with future repeat-surveys in mind once clearance of logged forest for oil palm plantation has resulted in the creation of forest fragments.
There are no areas of continuous, unfragmented old growth forest near to the SAFE project sites and hence the study design does not allow separation of the effects of location from those of habitat disturbance. We are therefore cautious in our interpretation of the results, particularly about assigning causal relationships between treatments and assemblage composition.
Survey work was conducted in April and May during the dry season, between h and h. None of the sites was affected by fire during the drought period, however.
Soil was removed from each pit and hand-searched for ants and termites using a white tray for 10 person-minutes. Bark was removed and holes in the wood were examined. These methods only sample the fauna living within the soil and dead wood, and do not sample the leaf litter community. Ants and termites were sorted to genus using the collections of the Natural History Museum, London, and relevant literature Ahmed and Akhtar ; Tho and Kirton ; Bolton ; Gathorne-Hardy ; Hashimoto Ant and termite reproductives were excluded from counts to avoid including vagrants, and immature termites could not be identified.
Ants and termites show niche conservatism within genera Andersen ; Donovan et al. Our approach is somewhat conservative, because species-rich genera, such as Pheidole and Strumigenys , are only counted as one occurrence per pit, despite being likely to be present as many species. Ants were assigned to functional groups following Andersen and Brown and termites to feeding groups following Donovan et al.
Ants were grouped according to differences in behaviour, dominance and temperature preferences in addition to feeding strategy, whereas termite groups were based only on feeding differences position along the humification gradient and associated morphological mandibular and gut structural characters Donovan et al. Differences in these ant and termite functional groups between treatments are therefore likely to be associated with differences in the rate of decomposition, the type of material being decomposed by termites and the extent and type of predation by ants.
The termite feeding group assignments represent the only widely-used functional classification system for this group. For ants, although morphological classifications Bihn et al.
Full details of genera within functional groups are listed in Tables 2 and 3. We measured the following environmental variables in each quadrat to assess habitat type and degree of disturbance: slope using a clinometer; percentage cover of leaf litter, bare ground, low vegetation, trees, dead wood, and grass following Cleary et al.
At each pit, leaf litter depth and humus depth were measured before digging. Humus depth was defined as depth mm of the dark, uppermost layer of soil between the decomposing leaf litter and lighter, more compact soil below.
Statistical analyses were conducted using R 2. Trends in genus richness and genus occurrence were consistent across soil and dead wood samples Online Resources, Table S2 , so data from both microhabitats were combined for use in all analyses.
We tested differences in both total and functional group occurrence across different habitat types using Kruskal—Wallis tests because occurrence data were not normally distributed and could not be normalised by transformation. Pairwise Wilcoxon rank sum tests with critical p -values reduced to account for multiple tests following Sokal and Rohlf , p were used to determine which habitats showed significant differences in occurrences.
Data on occurrence of ant and termite functional groups were first entered into a Detrended Correspondence Analysis DCA to assess gradient lengths. The significance of the association between each environmental variable with readings averaged for each quadrat and habitat type included as a dummy binary variables and variation in community functional structure were tested using Monte Carlo permutation tests with randomisations.
Forward selection was used to rank variables in order of importance in terms of their association with differences in species composition. This procedure selects the variable with the highest marginal eigenvalue followed, stepwise, by those with the highest eigenvalues conditional on the variance explained by all the previous steps Ter Braak and Verdonschot Both marginal effects explanatory effect of each variable when considered singly and conditional effects additional explanatory effect of each successive new variable when added by forward selection were calculated.
We chose to present the marginal effects rather than conditional effects since it cannot be assumed that the latter will select those variables with ecologically meaningful correlations with assemblage structure. Instead, displaying marginal effects allows a number of candidate explanatory variables to be visualised in relation to the major gradients of assemblage variation.
A total of 4, ants and 1, termites were sampled across soil pits and dead wood examinations. Encounters with ants were approximately three times more frequent than encounters with termites in old growth forest, 10 times more frequent in logged forest, and 25 times more frequent in oil palm plantation.
Functional group structure of both taxa varied with habitat type Fig. For the ants Fig. Mean occurrence of ants a and termites b per quadrat in old growth forest, logged forest and oil palm plantation. Shading indicates mean occurrence per group see legend. Termite feeding groups: Group I—feed on dead wood and grass; Group II—feed on grass, dead wood and leaf litter; Group IIF—feed on grass, dead wood and leaf litter with the help of fungal symbionts; Group III—feed on organic rich upper soil layers; Group IV—feed on organically poor soil.
See Online Resources, Table S3 for all statistical results. For ants, the strongest associations with functional group composition were: forest quality, humus depth, slope, cover of leaf litter, small saplings, grass and bare ground cover Table 4 a. These variables were included in the final RDA Fig.
Logged forest and grass cover were more strongly associated with axis 1 which largely comprises a gradient of occurrence of Tropical-climate Specialists and Subordinate Camponotini, both being found more commonly in logged forest with high grass cover Fig. The remaining significant environmental variables old growth forest, humus depth, leaf litter depth, forest quality, slope, small saplings cover, and bare ground cover were associated with axis 2 Fig.
In the latter case, all variables were positively associated, except for bare ground cover which was negatively associated. Ant functional groups were variable in their associations with this disturbance gradient Fig.
Ordination tri-plots showing redundancy analysis RDA of ant functional group occurrence a and termite feeding group occurrence b and marginally significant environmental variables in quadrats across all habitat types.
For ants a axis 1 explained For termites b axis 1 explained Abbreviations for functional and feeding groups are as for Fig. For termites, forest quality, slope, cover of tall poles, leaf litter and bare ground were strongly associated with feeding group structure Table 4 and were the variables included in the final RDA Fig. Old growth forest, forest quality, slope, tall poles and leaf litter cover were positively associated with axis 1, while logged forest and bare ground cover had negative axis 1 scores Fig.
Axis 1 of the RDA largely reflected associations with forest quality and a habitat disturbance gradient. All termite feeding groups were positively associated with axis 1 i. Axis 2 accounted for only 2. Group IIF and Group I showed stronger associations with axis 2 than axis 1, being positively and negatively associated with bare ground cover, respectively Fig. Worker termites and soldier termites are generally smaller.
Most termite species are found in the tropics, but several dozen species are found throughout the United States, including Hawaii, Puerto Rico, and the U. Virgin Islands. Alaska is the only U. Depending on the species, termite nests are found underground, in dead trees and stumps, in the tops of live trees, and in wooden structures. Ants are among the biggest predators of termites. Instead they have symbiotic protozoa and bacteria in their guts to break down the tough plant fibers.
Something all termite species share is a caste system. At the lowest level are the workers. These termites are sexually and developmentally immature. Workers take part in building out the nest, locating food, and caring for young. Soldiers are the next step up in the caste system. Like workers, they are immature in form, but in some species, they have well-developed mandibles for defense. Then there are the reproductives also called alates , winged males and females that are the product of mature colonies.
Alates swarm in warm weather and form pairs of males and females. There are several families of termites, and each has a different life history. Subterranean termites are found in both the eastern and western United States and cause the most damage to homes. The Formosan subterranean termite in the West is not native to the U.
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