In Peninsular Malaysia ten species of lycaenid butterflies use leaf flushes or inflorescences of the legume tree Saraca thaipingensis as larval hostplant. Resource partitioning among these species is regulated by a complex mixture of patterns of interaction with ants. Females of obligately myrmecophilous species lay their eggs exclusively on trees colonized by their specific host ants. On trees colonized by weaver ants, only specialist mutualists adapted to these territorial ants are able to survive, while larvae of other species are killed. The formicine ant Cladomyrma petalae, which inhabits hollow twigs of the myrmecophytic hostplant, likewise precludes oviposition by female butterflies. Lycaenid larvae confronted with this ant species never survive, but one concealed feeding species (Jamides caeruleus) escapes removal due to the cryptic life-habits of the larvae. Two facultative myrmecophiles associate in a mutualistic way with a wide and largely overlapping range of ant genera which forage at the extrafloral nectaries of leaf flushes. One species (Cheritra freja) is not myrmecophilous, but is tolerated by all but the most territorial ants. Ant-dependent hostplant selection and egg-clustering characterize the obligate mutualists, whereas facultative myrmecophiles and the non-myrmecophile distribute their eggs singly over appropriate hostplants. Signals mediating caterpillar-ant communication are highly specialized in one obligate myrmecophile (Drupadia theda), but rather unspecific in four other species tested. Altogether our observations indicate that colonization and establishment of lycaenid butterflies on S. thaipingensis trees are governed by specializations as well as opportunistic use of resources (ants and hostplant parts). Therefore, the diversity of this species assemblage is maintained by deterministic as well as stochastic factors.
1. Density of nests, population number, biomass, nitrogen and carbon content, calorific content and respiration rate of termites were studied at Pasoh Forest, West Malaysia mainly with four dominant species of epigeous nest builders, Macrotermes carbonarius, Dicuspiditermes nemorosus type-a, type-b and Homallotermes foraminifer, to reveal their role in the ecosystem. 2. The density of nests or mounds was 15-41/ha in M. carbonarius, 60-110/ha in the two types of D. nemorosus and 85-165/ha in H. foraminifer. 3. The population number per nest or mound was about 88,000 in M. carbonarius, 45,000 in D. nemorosus type-a, 47,000 in D. nemorosus type-b and 13,000 in H. foraminifer. The population number per hectare was about 1.8x10(6) for M. carbonarius, 4.3x10(6) for D. nemorosus type-a, 5.2x10(6) for D. nemorosus type-b and 2.1x10(6) for H. foraminifer. 4. The ratio in number of workers to soldiers was 6.4 for M. carbonarius, 19.0 for D. nemorosus type-a, 23.1 for D. nemorosus type-b and 8.9 for H. foraminifer. The ratio in number of adults (workers plus soldiers) to larvae was 1.4 for M. carbonarius, 0.59 for D. nemorosus type-a, 1.76 for D. nemorosus type-b and 3.84 for H. foraminifer. 5. The number of adults per nest of D. nemorosus type-a, type-b and H. foraminifer was linearly correlated with the weight of nest on the log-log coordinates. 6. The ratio of dry body weight to live weight was 0.24-0.30 for workers, 0.19-0.29 for soldiers and 0.15-0.22 for larvae. 7. The ash content of termite body was 26-66% for workers, 2-24% for soldiers and 1-4% for larvae. 8. The carbon content ranged from 45% to 66% of ash-free dry weight and the nitrogen content from 5.6% to 12.6%. 9. The mean calorific value of termite body was 5.3 gcal/mg in workers and soldiers on an ash-free weight basis, but was greater in nymphs and winged reproductives in the nest (6.7-6.9 gcal/mg) owing probably to their large fat storage. 10. The relation of CO2 evolution rate to temperature in these termites was similar to what has been found in other insects. 11. The total biomass of the four termite species was estimated at 6.01 kg ash-free dry weight/ha, equivalent to 0.55 kg nitrogen/ha and 3.09 kg carbon/ha. 12. The role of the fungi cultivated by M. carbonarius on their fungus combs was discussed in relation to the nutrition of termites and the decomposition of leaf-litter with special reference to their nitrogen metabolism. The high nitrogen content of fungus spherules growing on fungus combs seemed to have an important bearing on the nutrition of termites. 13. It was concluded that the termites played a very important role in the organic matter decomposition cycle of Pasoh Forest.
To estimate the rate of consumption of leaf litter by termites on the forest floor of Pasoh Forest Reserve, Negeri Sembilan, West Malaysia, newly fallen leaves were marked and distributed on the ground. The loss of leaf area due to termites was determined either photometrically or visually. An average of 1.70% of the total surface area of the leaf litter disappeared per week in experiment 1 and 1.25% in experiment 2 in Plot 1, and 2.9% per day in other plots located near the mounds of Macrotermes carbonarius. The amount of leaf litter accumulation in the Ao layer was estimated at about 2.3 t/ha at Plot 1, so it was likely that an amount equivalent to about 32% of the daily leaf-litter fall was transported by M. carbonarius to their mounds in experiment 1 and 22% in experiment 2. It was considered that the termites had an important role in the detritus food chain of the ecosystem.
Tropical rainforest disturbance and conversion are critical drivers of biodiversity loss. A key knowledge gap is understanding the impacts of habitat modification on mechanisms of community assembly, which are predicted to respond differently between taxa and across spatial scales. We use a null model approach to detect trait assembly of species at local- and landscape-scales, and then subdivide communities with different habitat associations and foraging guilds to investigate whether the detection of assembly mechanisms varies between groups. We focus on two indicator taxa, dung beetles and birds, across a disturbance gradient of primary rainforest, selectively logged rainforest, and oil palm plantations in Borneo, Southeast Asia. Random community assembly was predominant for dung beetles across habitats, whereas trait convergence, indicative of environmental filtering, occurred across the disturbance gradient for birds. Assembly patterns at the two spatial scales were similar. Subdividing for habitat association and foraging guild revealed patterns hidden when focusing on the overall community. Dung beetle forest specialists and habitat generalists showed opposing assembly mechanisms in primary forest, community assembly of habitat generalists for both taxa differed with disturbance intensity, and insectivorous birds strongly influenced overall community assembly relative to other guilds. Our study reveals the sensitivity of community assembly mechanisms to anthropogenic disturbance via a shift in the relative contribution of stochastic and deterministic processes. This highlights the need for greater understanding of how habitat modification alters species interactions and the importance of incorporating species' traits within assessments.