We investigate the geographical and historical context of diversification in a complex of mutualistic Crematogaster ants living in Macaranga trees in the equatorial rain forests of Southeast Asia. Using mitochondrial DNA from 433 ant colonies collected from 32 locations spanning Borneo, Malaya and Sumatra, we infer branching relationships, patterns of genetic diversity and population history. We reconstruct a time frame for the ants' diversification and demographic expansions, and identify areas that might have been refugia or centres of diversification. Seventeen operational lineages are identified, most of which can be distinguished by host preference and geographical range. The ants first diversified 16-20 Ma, not long after the onset of the everwet forests in Sundaland, and achieved most of their taxonomic diversity during the Pliocene. Pleistocene demographic expansions are inferred for several of the younger lineages. Phylogenetic relationships suggest a Bornean cradle and major axis of diversification. Taxonomic diversity tends to be associated with mountain ranges; in Borneo, it is greatest in the Crocker Range of Sabah and concentrated also in other parts of the northern northwest coast. Within-lineage genetic diversity in Malaya and Sumatra tends to also coincide with mountain ranges. A series of disjunct and restricted distributions spanning northern northwest Borneo and the major mountain ranges of Malaya and Sumatra, seen in three pairs of sister lineages, further suggests that these regions were rain-forest refuges during drier climatic phases of the Pleistocene. Results are discussed in the context of the history of Sundaland's rain forests.
Gnamptogenys menadensis is an arboreal nester that forages opportunistically almost exclusively on vegetation, sometimes recruiting others to participate in prey retrieval. The three-dimensional characteristics of vegetation suggest that functions describing recruitment decision thresholds or the pattern of recruitment in arboreal species may differ from those predicted by optimal foraging theory. To examine the effects of prey abundance and distance on the recruitment dynamics of G. menadensis, we baited nests with one termite, five termites or a number of termites between 20 and 40 either near to or far from the entrance and observed the ensuing behaviors. G. menadensis recruited others when encountering multiple termites regardless of the termite pile's distance from the nest, although a few individuals remained at the site and defended the resource. The pattern of arrivals at the site indicates that the majority and sometimes all arrivals were recruited from the branch trails. In combination, these results suggest that the architecture of the foraging habitat, which limits available return routes to the nest and thus increases encounter probabilities with potential recruits, shaped the process of information transfer and generated a collective pattern of foraging and prey retrieval.
South East Asia is widely regarded as a centre of threatened biodiversity owing to extensive logging and forest conversion to agriculture. In particular, forests degraded by repeated rounds of intensive logging are viewed as having little conservation value and are afforded meagre protection from conversion to oil palm. Here, we determine the biological value of such heavily degraded forests by comparing leaf-litter ant communities in unlogged (natural) and twice-logged forests in Sabah, Borneo. We accounted for impacts of logging on habitat heterogeneity by comparing species richness and composition at four nested spatial scales, and examining how species richness was partitioned across the landscape in each habitat. We found that twice-logged forest had fewer species occurrences, lower species richness at small spatial scales and altered species composition compared with natural forests. However, over 80 per cent of species found in unlogged forest were detected within twice-logged forest. Moreover, greater species turnover among sites in twice-logged forest resulted in identical species richness between habitats at the largest spatial scale. While two intensive logging cycles have negative impacts on ant communities, these degraded forests clearly provide important habitat for numerous species and preventing their conversion to oil palm and other crops should be a conservation priority.
The mechanism by which palatable species take advantage of their similarity in appearance to those that are unpalatable, in order to avoid predation, is called Batesian mimicry. Several arthropods are thought to be Batesian mimics of social insects; however, social insects that are Batesian mimics among themselves are rare. In Malaysia we found a possible Batesian mimic in an arboreal ant species, Camponotus sp., which was exclusively observed on foraging trails of the myrmicine ant Crematogaster inflata. The bright yellow and black colouring pattern, as well as the walking behaviour, were very similar in both species. We observed general interactions between the two species, and tested their palatability and the significance of the remarkably similar visual colour patterns for predator avoidance. Prey offered to C. inflata was also eaten by Camponotus workers in spite of their being attacked by C. inflata, indicating that Camponotus sp. is a commensal of C. inflata. An experiment with chicks as potential predators suggests that Camponotus sp. is palatable whereas C. inflata is unpalatable. After tasting C. inflata, the chicks no longer attacked Camponotus sp., indicating that Camponotus sp. is a Batesian mimic of Crematogaster inflata.
A key driver of rain forest degradation is rampant commercial logging. Reduced-impact logging (RIL) techniques dramatically reduce residual damage to vegetation and soils, and they enhance the long-term economic viability of timber operations when compared to conventionally managed logging enterprises. Consequently, the application of RIL is increasing across the tropics, yet our knowledge of the potential for RIL also to reduce the negative impacts of logging on biodiversity is minimal. We compare the impacts of RIL on birds, leaf-litter ants, and dung beetles during a second logging rotation in Sabah, Borneo, with the impacts of conventional logging (CL) as well as with primary (unlogged) forest. Our study took place 1-8 years after the cessation of logging. The species richness and composition of RIL vs. CL forests were very similar for each taxonomic group. Both RIL and CL differed significantly from unlogged forests in terms of bird and ant species composition (although both retained a large number of the species found in unlogged forests), whereas the composition of dung beetle communities did not differ significantly among forest types. Our results show little difference in biodiversity between RIL and CL over the short-term. However, biodiversity benefits from RIL may accrue over longer time periods after the cessation of logging. We highlight a severe lack of studies investigating this possibility. Moreover, if RIL increases the economic value of selectively logged forests (e.g., via REDD+, a United Nations program: Reducing Emissions from Deforestation and Forest Degradation in Developing Countries), it could help prevent them from being converted to agricultural plantations, which results in a tremendous loss of biodiversity.
Rattan is an important climbing palm taxon in Malaysian tropical rain forests. Many rattan species have unique structures directly associated with certain ant species. In this study, four rattan species (Daemonorops lewisiana, Calamus castaneus, Daemonorops geniculata and Korthalsia scortechinii) were inspected and documented in a field survey concerning their relationships with several ant species. We noticed that two rattan species (D. lewisiana and C. castaneus) were more likely to be associated with ants compared to their neighbouring rattan (Plectomia griffithii). However, D. lewisiana and C. castaneus did not directly provide shelters for ant colonies, but possessed unique structures: upward-pointing spines and funnel-shaped leaves, which are equipped to collect more litter than P. griffithii. To test our litter collecting hypothesis, we measured the inclination of spines from the stem. Our results showed the presence of ant colonies in the litter-collecting rattans (D. lewisiana and C. castaneus), which was significantly higher compared to a non-litter-collecting rattan (P. griffithii). We propose a complex and novel type of adaptation (litter-collection and provision of nesting materials) for rattans, which promotes interactions between the rattan and ants through the arrangements of leaves, leaflets, and spines. In return, the rattan may benefit from ants' services, such as protection, nutrient enhancement, and pollination.
Invasive species are one of the main sources of the ongoing global loss of biodiversity. Invasive ants are known as particularly damaging invaders and their introductions are often accompanied by population-level behavioural and genetic changes that may contribute to their success. Anoplolepis gracilipes is an invasive ant that has just recently received increased attention due to its negative impact on native ecosystems. We examined the behaviour and population structure of A. gracilipes in Sabah, Malaysia. A total of 475 individuals from 24 colonies were genotyped with eight microsatellite markers. Intracolonial relatedness was high, ranging from 0.37 to 1 (mean +/- SD: 0.82 +/- 0.04), while intercolonial relatedness was low (0.0 +/- 0.02, range -0.5-0.76). We compared five distinct sampling regions in Sabah and Brunei. A three-level hierarchical F-analysis revealed high genetic differentiation among colonies within the same region, but low genetic differentiation within colonies or across regions. Overall levels of heterozygosity were unusually high (mean H(O) = 0.95, mean H(E) = 0.71) with two loci being entirely heterozygous, indicating an unusual reproductive system in this species. Bioassays revealed a negative correlation between relatedness and aggression, suggesting kinship as one factor facilitating supercolony formation in this species. Furthermore, we genotyped one individual per nest from Sabah (22 nests), Sarawak (one nest), Brunei (three nests) and the Philippines (two nests) using two mitochondrial DNA markers. We found six haplotypes, two of which included 82.1% of all sequences. Our study shows that the sampled area in Sabah consists of a mosaic of differently interrelated nests in different stages of colony establishment. While some of the sampled colonies may belong to large supercolonies, others are more likely to represent recently introduced or dispersed propagules that are just beginning to expand.
The longlegged ant, Anoplolepis gracilipes (Fr. Smith) (Hymenoptera: Formicidae), is a highly invasive species that can aggressively displace other ant species. We conducted laboratory assays to examine interspecies aggression of A. gracilipes versus 15 sympatric ant species found in the urban environment and disturbed habitat in Malaysia: Monomorium pharaonis (L.), Monomorium floricola (Jerdon), Monomorium orientale Mayr, Monomorium destructor (Jerdon), Pheidole parva Mayr, Crematogaster sp., Solenopsis geminata (F.), Tapinoma indicum (Forel), Tapinoma melanocephalum (F.), Technomyrmnex butteli Forel, Dolichoderus thoracicus (Smith), Paratrechina longicornis (Latrielle), Oecophylla smaragdina (F), Camponotus sp., and Tetraponera rufonigra (Jerdon). A. gracilipes showed aggressive behavior toward all opponent species, except the smallest M. orientale. Opponent species size (body size, head width, and mandible width) was significantly correlated with A. gracilipes aggression level and mortality rate. We also found a significant positive relationship between A. gracilipes aggression level and the mortality of the opponent species. The results suggest that invasive populations of A. gracilipes would have the greatest impact on larger ant species. In addition, we examined the intraspecific aggression of A. gracilipes. We found that A. gracilipes from different localities in Malaysia showed intraspecific aggression toward one another. This finding differs from the results of studies conducted in Christmas Island earlier. Differences in the genetic variability among populations may explain these differing results.
Budding and relocation of nests are important characteristics of the Pharaoh ant, Monomorium pharaonis (L.), an important pest of artificial structures. Pharaoh ant colony movements induced by several types of disturbances were evaluated in the laboratory. The percentages of workers and brood in the source and new nest sites were determined at Days 0, 1, 3, and 5 following physical disturbance (temporal removal of nestmates), chemical disturbance (application of pyrethroid insecticide), invasion by heterospecific ants, food depletion, and moisture depletion in the laboratory. All disturbances were performed in the source nest, which was connected to an empty new nest site. Almost all workers moved and carried the entire brood to the new nest site when subjected to physical disturbance, chemical disturbance, and ant invasion on Day 1, whereas only <5% of workers were present in the new nest site in the undisturbed control. After these disturbances, the brood was never relocated back to the original nest site in this 5-d study. When subjected to food depletion, ∼60% of the brood were found in the new nest site and ∼40% of the brood remained in the original nest on Day 5, resulting in a polydomous population. In contrast, moisture depletion did not show any significant effect on colony movement. These results provide useful information about the causes of Pharaoh ant colony budding and guidance about how to develop effective control and prevention strategies.
Trophic organisation defines the flow of energy through ecosystems and is a key component of community structure. Widespread and intensifying anthropogenic disturbance threatens to disrupt trophic organisation by altering species composition and relative abundances and by driving shifts in the trophic ecology of species that persist in disturbed ecosystems. We examined how intensive disturbance caused by selective logging affects trophic organisation in the biodiversity hotspot of Sabah, Borneo. Using stable nitrogen isotopes, we quantified the positions in the food web of 159 leaf-litter ant species in unlogged and logged rainforest and tested four predictions: (i) there is a negative relationship between the trophic position of a species in unlogged forest and its change in abundance following logging, (ii) the trophic positions of species are altered by logging, (iii) disturbance alters the frequency distribution of trophic positions within the ant assemblage, and (iv) disturbance reduces food chain length. We found that ant abundance was 30% lower in logged forest than in unlogged forest but changes in abundance of individual species were not related to trophic position, providing no support for prediction (i). However, trophic positions of individual species were significantly higher in logged forest, supporting prediction (ii). Consequently, the frequency distribution of trophic positions differed significantly between unlogged and logged forest, supporting prediction (iii), and food chains were 0.2 trophic levels longer in logged forest, the opposite of prediction (iv). Our results demonstrate that disturbance can alter trophic organisation even without trophically-biased changes in community composition. Nonetheless, the absence of any reduction in food chain length in logged forest suggests that species-rich arthropod food webs do not experience trophic downgrading or a related collapse in trophic organisation despite the disturbance caused by logging. These food webs appear able to bend without breaking in the face of some forms of anthropogenic disturbance.
Monomorium orientale Mayr (Hymenoptera: Formicidae) is a common structure- and food-infesting ant in Asia. There is only limited information on the biology and habits of this species, especially on the preferred foods and distribution of nutrients in colonies. We conducted a laboratory study on the distribution of carbohydrates, proteins, and lipids, which were represented by respective food sources, in M. orientale colonies. Three colony conditions were applied: normal, with a balanced ratio of castes, queenless (only workers and brood), and broodless (only queens and workers). Food sources were stained to track the flow of the respective food in the colonies. Results revealed that carbohydrates had rapid distribution, with > 60% of the colony indicated in 24 h, in all colony conditions. Queens in all colonies did not feed on protein. Protein showed a more delayed distribution in the brood in all colony conditions; < 10% of the colony fed on protein by 24 h. Only queens in broodless colonies showed signs of feeding on lipid, with < 10% indicated in 24 h. Workers in all colonies fed on lipid as soon as it was delivered, whereas the brood only began to reveal feeding response after 24 h.
Understanding how anthropogenic disturbance influences patterns of community composition and the reinforcing interactive processes that structure communities is important to mitigate threats to biodiversity. Competition is considered a primary reinforcing process, yet little is known concerning disturbance effects on competitive interaction networks. We examined how differences in ant community composition between undisturbed and disturbed Bornean rainforest, is potentially reflected by changes in competitive interactions over a food resource. Comparing 10 primary forest sites to 10 in selectively-logged forest, we found higher genus richness and diversity in the primary forest, with 18.5% and 13.0% of genera endemic to primary and logged respectively. From 180 hours of filming bait cards, we assessed ant-ant interactions, finding that despite considered aggression over food sources, the majority of ant interactions were neutral. Proportion of competitive interactions at bait cards did not differ between forest type, however, the rate and per capita number of competitive interactions was significantly lower in logged forest. Furthermore, the majority of genera showed large changes in aggression-score with often inverse relationships to their occupancy rank. This provides evidence of a shuffled competitive network, and these unexpected changes in aggressive relationships could be considered a type of competitive network re-wiring after disturbance.
Termites and ants contribute more to animal biomass in tropical rain forests than any other single group and perform vital ecosystem functions. Although ants prey on termites, at the community level the linkage between these groups is poorly understood. Thus, assessing the distribution and specificity of ant termitophagy is of considerable interest. We describe an approach for quantifying ant-termite food webs by sequencing termite DNA (cytochrome c oxidase subunit II, COII) from ant guts and apply this to a soil-dwelling ant community from tropical rain forest in Gabon. We extracted DNA from 215 ants from 15 species. Of these, 17.2 % of individuals had termite DNA in their guts, with BLAST analysis confirming the identity of 34.1 % of these termites to family level or better. Although ant species varied in detection of termite DNA, ranging from 63 % (5/7; Camponotus sp. 1) to 0 % (0/7; Ponera sp. 1), there was no evidence (with small sample sizes) for heterogeneity in termite consumption across ant taxa, and no evidence for species-specific ant-termite predation. In all three ant species with identifiable termite DNA in multiple individuals, multiple termite species were represented. Furthermore, the two termite species that were detected on multiple occasions in ant guts were in both cases found in multiple ant species, suggesting that ant-termite food webs are not strongly compartmentalised. However, two ant species were found to consume only Anoplotermes-group termites, indicating possible predatory specialisation at a higher taxonomic level. Using a laboratory feeding test, we were able to detect termite COII sequences in ant guts up to 2 h after feeding, indicating that our method only detects recent feeding events. Our data provide tentative support for the hypothesis that unspecialised termite predation by ants is widespread and highlight the use of molecular approaches for future studies of ant-termite food webs.
A field study on foraging activity and proteinacous food preference was performed on the tropical fire ant (Solenopsis geminata) (Fabricius) at the School of Biological Sciences and Desasiswa Bakti Permai, Universiti Sains Malaysia (USM), Penang. Foraging activity studies of 4 colonies of S. geminata were conducted in the field for 24 hours. Foraging activity significantly increased 4 hours before sunset and maximum foraging occurred at midnight until early morning. Three types of proteinacous food; anchovy, meat and egg yolk were tested among the five colonies of S. geminata in the field. The egg yolk was the most preferred food (100%) followed by meat (31%) and anchovy (15%).
The behavioral response of the obligate bamboo-nesting ant Cataulacus muticus to nest flooding was studied in a perhumid tropical rainforest in Malaysia and in the laboratory. The hollow internodes of giant bamboo, in which C. muticus exclusively nests, are prone to flooding by heavy rains. The ants showed a two-graded response to flooding. During heavy rain workers block the nest entrances with their heads to reduce water influx. However, rainwater may still intrude into the nest chamber. The ants respond by drinking the water, leaving the nest and excreting water droplets on the outer stem surface. This cooperative 'peeing' behavior is a new survival mechanism adaptive to the ants' nesting ecology. Laboratory experiments conducted with two other Cataulacus species, C. catuvolcus colonizing small dead twigs and C. horridus inhabiting rotten wood, did not reveal any form of water-bailing behavior.
We investigate the evolution of host association in a cryptic complex of mutualistic Crematogaster (Decacrema) ants that inhabits and defends Macaranga trees in Southeast Asia. Previous phylogenetic studies based on limited samplings of Decacrema present conflicting reconstructions of the evolutionary history of the association, inferring both cospeciation and the predominance of host shifts. We use cytochrome oxidase I (COI) to reconstruct phylogenetic relationships in a comprehensive sampling of the Decacrema inhabitants of Macaranga. Using a published Macaranga phylogeny, we test whether the ants and plants have cospeciated. The COI phylogeny reveals 10 well-supported lineages and an absence of cospeciation. Host shifts, however, have been constrained by stem traits that are themselves correlated with Macaranga phylogeny. Earlier lineages of Decacrema exclusively inhabit waxy stems, a basal state in the Pachystemon clade within Macaranga, whereas younger species of Pachystemon, characterized by nonwaxy stems, are inhabited only by younger lineages of Decacrema. Despite the absence of cospeciation, the correlated succession of stem texture in both phylogenies suggests that Decacrema and Pachystemon have diversified in association, or codiversified. Subsequent to the colonization of the Pachystemon clade, Decacrema expanded onto a second clade within Macaranga, inducing the development of myrmecophytism in the Pruinosae group. Confinement to the aseasonal wet climate zone of western Malesia suggests myrmecophytic Macaranga are no older than the wet forest community in Southeast Asia, estimated to be about 20 million years old (early Miocene). Our calculation of COI divergence rates from several published arthropod studies that relied on tenable calibrations indicates a generally conserved rate of approximately 1.5% per million years. Applying this rate to a rate-smoothed Bayesian chronogram of the ants, the Decacrema from Macaranga are inferred to be at least 12 million years old (mid-Miocene). However, using the extremes of rate variation in COI produces an age as recent as 6 million years. Our inferred timeline based on 1.5% per million years concurs with independent biogeographical events in the region reconstructed from palynological data, thus suggesting that the evolutionary histories of Decacrema and their Pachystemon hosts have been contemporaneous since the mid-Miocene. The evolution of myrmecophytism enabled Macaranga to radiate into enemy-free space, while the ants' diversification has been shaped by stem traits, host specialization, and geographic factors. We discuss the possibility that the ancient and exclusive association between Decacrema and Macaranga was facilitated by an impoverished diversity of myrmecophytes and phytoecious (obligately plant inhabiting) ants in the region.
Tarsal adhesive pads enable insects to hold on to smooth plant surfaces. Using a centrifuge technique, we tested whether a "wet adhesion" model of a thin film of liquid secreted between the pad and the surface can explain adhesive and frictional forces in Asian Weaver ants (Oecophylla smaragdina). When forces are acting parallel to the surface, pads in contact with the surface can slide smoothly. Force per unit pad contact area was strongly dependent on sliding velocity and temperature. Seemingly consistent with the effect of a thin liquid film in the contact zone, (1) frictional force linearly increased with sliding velocity, (2) the increment was greater at lower temperatures and (3) no temperature dependence was detected for low-rate perpendicular detachment forces. However, we observed a strong, temperature-independent static friction that was inconsistent with a fully lubricated contact. Static friction was too large to be explained by the contribution of other (sclerotized) body parts. Moreover, the rate-specific increase of shear stress strongly exceeded predictions derived from estimates of the adhesive liquid film's thickness and viscosity. Both lines of evidence indicate that the adhesive secretion alone is insufficient to explain the observed forces and that direct interaction of the soft pad cuticle with the surface ("rubber friction") is involved.
Ants are social insects with some significant roles in the ecosystem, including acting as predators for various insect pests. Myopopone castanea ants is a predatorfor the larvae of Oryctes rhinoceros pest. The existence of a similar niche of life between M. castanea ants and O. rhinoceros larvae opens an excellent opportunity to utilize these ants as biological agents. The research was conducted to study some aspects biology of M. castanea so that later it can be applied to mass rearing of natural enemies in the laboratory. The study was conducted by maintaining 50 eggs of M. castanea ant. Then, the eggs placed on two pieces of decayed palm oil stem together with twenty individual worker ants and ten individual end instar larvae. It needs five replications for the experiment. The results showed that egg stadia length was 13.8 days. It found five instars within M. castanea ant larvae with varying lengths of each stage. It takes 17.2 days for worker ant pupae to go through stadia pupa and 17.9 days for female ant pupae. The survival rate of M. castanea ant life from eggs until imago is 56.4%, which means that from several groups of eggs laid by queen ants, only about half have succeeded in becoming ant imago.