Inter-specific aggression generates ant mosaics in canopies of primary tropical rainforest

The ant mosaic is a concept of the non-random spatial distribution of individual ant species in trees built upon the assumption of interspecific behavioural associations. However, colony identity and environmental variance may also play a role in species distribution. Here we assess the presence of ant mosaics in a primary forest ecosystem and whether they are structured by species' aggressive behaviours or by habitat filtering. We sampled arboreal ants from vertically stratified baits exposed in 225 canopy trees in a 9-ha plot of primary lowland forest in Papua New Guinea, the largest forest area surveyed to detect ant mosaics. We performed behavioural tests on conspecific ants from adjacent trees to determine the territories of individual colonies. We explored the environmental effects on the ant communities using information on the plot vegetation structure and topography. Furthermore, we created a novel statistical method to test for the community non-random spatial structure across the plot via spatial randomisation of individual colony territories. Finally, we linked spatial segregation among the four most common species to experimentally assessed rates of interspecies aggression. The ant communities comprised 57 species of highly variable abundance and vertical stratification. Ant community composition was spatially dependent, but it was not affected by tree species composition or canopy connectivity. Only local elevation had a significant but rather small effect. Individual colony territories ranged from one tree to 0.7 ha. Species were significantly over-dispersed, with their territory overlap significantly reduced. The level of aggression between pairs of the four most common species was positively correlated with their spatial segregation. Our study demonstrates the presence of ant mosaics in tropical pristine forest, which are maintained by interspecific aggression rather than habitat filtering, with vegetation structure having a rather small and indirect effect, probably linked to microclimate variability.     

Bottom-up control and co-occurrence in complex communities: honeydew and nectar determine a rainforest ant mosaic

Complex distribution patterns of species-rich insect communities in tropical rainforests have been intensively studied, and yet we know very little about processes that generate these patterns. We provide evidence for the key role of homopteran honeydew and plant nectar in structuring ant communities in an Australian tropical rainforest canopy and understorey. We also test the ant visitation of these resources against predictions derived from the 'ant-mosaic' hypothesis. Two ant species were highly dominant in terms of territorial behaviour and abundance: Oecophylla smaragdina and Anonychomyrma gilberti . Both dominant ant species monopolised large aggregations of honeydew-producing homopterans. Attended homopteran species were highly segregated between these two ant species. For the use of extrafloral and floral nectar (involving 43 ant species on 48 plant species), partitioning of ant species among plant species and between canopy and understorey was also significant, but less pronounced. In contrast to trophobioses, simultaneous co-occurrence of different nectar foraging ant species on the same plant individuals was frequent (23% of all surveys). While both dominant ant species were mutually exclusive on honeydew and nectar sources, co-occurrence with non-dominant ant species on nectaries was common. The proportion of visits with co-occurrences was low for dominant ants and high for many sub-ordinate species. These findings support the ant mosaic theory. The differential role of honeydew (as a specialised resource for dominant ants) and nectar (as an opportunistic resource for all ants including the co-occurring non-dominant species) provides a plausible structuring mechanism for the Australian canopy ant community studied.     

The dynamics of ant mosaics in tropical rainforests characterized using the Self‐Organizing Map algorithm

Ants, the most abundant taxa among canopy‐dwelling animals in tropical rainforests, are mostly represented by territorially dominant arboreal ants (TDAs) whose territories are distributed in a mosaic pattern (arboreal ant mosaics). Large TDA colonies regulate insect herbivores, with implications for forestry and agronomy. What generates these mosaics in vegetal formations, which are dynamic, still needs to be better understood. So, from empirical research based on 3 Cameroonian tree species (Lophira alata, Ochnaceae; Anthocleista vogelii, Gentianaceae; and Barteria fistulosa, Passifloraceae), we used the Self‐Organizing Map (SOM, neural network) to illustrate the succession of TDAs as their host trees grow and age. The SOM separated the trees by species and by size for L. alata, which can reach 60 m in height and live several centuries. An ontogenic succession of TDAs from sapling to mature trees is shown, and some ecological traits are highlighted for certain TDAs. Also, because the SOM permits the analysis of data with many zeroes with no effect of outliers on the overall scatterplot distributions, we obtained ecological information on rare species. Finally, the SOM permitted us to show that functional groups cannot be selected at the genus level as congeneric species can have very different ecological niches, something particularly true for Crematogaster spp., which include a species specifically associated with B. fistulosa, nondominant species and TDAs. Therefore, the SOM permitted the complex relationships between TDAs and their growing host trees to be analyzed, while also providing new information on the ecological traits of the ant species involved.     

Ant mosaics occur in SE Asian oil palm plantation but not rain forest and are influenced by the presence of nest‐sites and non‐native species

Interaction networks within biotic communities can be dramatically altered by anthropogenic habitat modification. Ants, an important ecological group, often interact competitively to form mosaic‐like patterns in disturbed plantation habitats, in which dominant species form mutually exclusive territories. However, the existence of these ant mosaics in pristine forests is contentious. Here we assess the relative strengths of ant competitive interactions in oil palm plantation and primary rain forest in Sabah, Malaysia, using null models of species co‐occurrence. We use two metrics: the C‐score, which measures mean degree of overall co‐occurrence, and a novel metric, the C_var‐score, which measures the variance in degree of co‐occurrence. We also investigate the role of nest sites by collecting ants from canopy and leaf litter microhabitats, and from epiphytic ferns, an important nest site for canopy ants. Furthermore, we assess whether non‐native species, which were widespread in oil palm plantation (61 occurrences vs five in rain forest) are important in driving the formation of ant mosaics. We found no evidence for ant mosaics in any primary forest microhabitat. In oil palm plantation, segregation between species was pronounced in epiphytes, weak in the rest of the canopy and absent in leaf litter communities. Intriguingly, exclusion of non‐native ant species from analyses increased the degree of negative species co‐occurrence in all three microhabitats, with species segregation in the oil palm canopy becoming statistically significant. Our results suggest that invasion of plantation habitats by non‐native species does not drive increased species segregation in ant communities. Rather, high degrees of species segregation might relate to changes in the importance of canopy nest sites, with colonies competing more strongly for these in plantations. In primary forests, weaker nest‐site limitation and the highly complex, more vertically stratified, non‐uniform canopy could lead to random co‐occurrence between ant species at the scales studied here.     

The arboreal ants of a Neotropical rain forest show high species density and comprise one third of the ant fauna

In tropical rain forests, the ant community can be divided into ground and arboreal faunas. Here, we report a thorough sampling of the arboreal ant fauna of La Selva Biological Station, a Neotropical rain forest site. Forty-five canopy fogging samples were centered around large trees. Individual samples harbored an average of 35 ant species, with up to 55 species in a single sample. The fogging samples yielded 163 observed species total, out of a statistically estimated 199 species. We found no relationship between within-sample ant richness and focal tree species, nor were the ant faunas of nearby trees more similar to each other than the faunas of widely spaced trees. Species density was high, and beta diversity was low: A single column of vegetation typically harbors at least a fifth of the entire arboreal ant fauna. Considering the entire fauna, based on 23,326 species occurrence records using a wide variety of collecting methods, 182 of 539 observed species (196 of 605, estimated statistically) were entirely arboreal. The arboreal ant fauna is thus about a third of the total La Selva ant fauna, a robust result because inventory completeness was similar for ground and arboreal ants. The taxonomic history of discovery of the species that make up the La Selva fauna reveals no disproportionately large pool of undiscovered ant species in the canopy. The "last biotic frontier" for tropical ants has been the rotten wood, leaf litter, and soil of the forest floor.     

Effects of selective logging on the arboreal ants of a Bornean rainforest

We examined the effect of selective logging and corresponding forest canopy loss on arboreal ant diversity in a tropical rainforest. Arboreal ants were collected from an unlogged forest plot and from forest plots selectively logged 14 years and 24 years earlier in Danum Valley, Sabah, Malaysia, using a canopy fogging method. Selective logging was associated with a significant decrease in canopy cover and an increase in understory vegetation density relative to unlogged forest. Our study showed that selective logging in primary forest might not dramatically decrease total species number and overall abundance of arboreal ants; however, it may influence the species composition and dominance structure of the ant community, accompanied by an increase in abundance of shrub‐layer species and trophobiotic species. In view of the results of this study, management techniques that minimize logging impact on understory vegetation structure are likely to help maintain the conservation value of logged forests for arboreal ants. Our results also suggest that accurate assessment of the impacts on biodiversity should not be based only on measurement of species number and overall abundance, but also on analysis of species composition and community structure.     

Biotic and abiotic determinants of the formation of ant mosaics in primary Neotropical rainforests

1. Ants are widespread in tropical rainforests, including in the canopy where territorially dominant arboreal species represent the main part of the arthropod biomass. 2. By mapping the territories of dominant arboreal ant species and using a null model analysis and a pairwise approach this study was able to show the presence of an ant mosaic on the upper canopy of a primary Neotropical rainforest (c. 1 ha sampled; 157 tall trees from 28 families). Although Neotropical rainforest canopies are frequently irregular, with tree crowns at different heights breaking the continuity of the territories of dominant ants, the latter are preserved via underground galleries or trails laid on the ground. 3. The distribution of the trees influences the structure of the ant mosaic, something related to the attractiveness of tree taxa for certain arboreal ant species rather than others. 4. Small‐scale natural disturbances, most likely strong winds in the area studied (presence of canopy gaps), play a role by favouring the presence of two ant species typical of secondary formations: Camponotus femoratus and Crematogaster levior, which live in parabiosis (i.e. share territories and nests but lodge in different cavities) and build conspicuous ant gardens. In addition, pioneer Cecropia myrmecophytic trees were recorded.     

An Ant Mosaic Revisited: Dominant Ant Species Disassemble Arboreal Ant Communities but Co-Occur Randomly

The spatial distributions of many tropical arboreal ant species are often arranged in a mosaic such that dominant species have mutually exclusive distributions among trees. These dominant species can also mediate the structure of the rest of the arboreal ant community. Little attention has been paid to how diet might shape the effects of dominant species on one another and the rest of the ant community. Here, we take advantage of new information on the diets of many tropical arboreal ant species to examine the intra- and inter-guild effects of dominant species on the spatial distribution of one another and the rest of the tropical arboreal ant community in a cocoa farm in Bahia, Brazil. Using null model analyses, we found that all ant species, regardless of dominance status or guild membership, co-occur much less than expected by chance. Surprisingly, the suite of five dominant species showed random co-occurrence patterns, suggesting that interspecific competition did not shape their distribution among cocoa trees. Across all species, there was no evidence that competition shaped co-occurrence patterns within guilds. Co-occurrence patterns of subordinant species were random on trees with dominant species, but highly nonrandom on trees without dominant species, suggesting that dominant species disassemble tropical arboreal ant communities. Taken together, our results highlight the often complex nature of interactions that structure species-rich tropical arboreal ant assemblages.     

Within-tree distribution of nest sites and foraging areas of ants on canopy trees in a tropical rainforest in Borneo

It has been argued that canopy trees in tropical rainforests harbor species-rich ant assemblages; however, how ants partition the space on trees has not been adequately elucidated. Therefore, we investigated within-tree distributions of nest sites and foraging areas of individual ant colonies on canopy trees in a tropical lowland rainforest in Southeast Asia. The species diversity and colony abundance of ants were both significantly greater in crowns than on trunks. The concentration of ant species and colonies in the tree crown seemed to be associated with greater variation in nest cavity type in the crown, compared to the trunk. For ants nesting on canopy trees, the numbers of colonies and species were both higher for ants foraging only during the daytime than for those foraging at night. Similarly, for ants foraging on canopy trees, both values were higher for ants foraging only during the daytime than for those foraging at night. For most ant colonies nesting on canopy trees, foraging areas were limited to nearby nests and within the same type of microhabitat (within-tree position). All ants foraging on canopy trees in the daytime nested on canopy trees, whereas some ants foraging on the canopy trees at night nested on the ground. These results suggest that spatial partitioning by ant assemblages on canopy trees in tropical rainforests is affected by microenvironmental heterogeneity generated by three-dimensional structures (e.g., trees, epiphytes, lianas, and aerial soils) in the crowns of canopy trees. Furthermore, ant diversity appears to be enriched by both temporal (diel) and fine-scale spatial partitioning of foraging activity.     

热带森林不同演替阶段蚂蚁巢穴的分布特征及其影响因素

为探明热带森林蚂蚁巢穴的分布特征及其影响因素, 采用样方法研究了西双版纳不同演替阶段热带森林定居巢穴蚂蚁的种类及其巢穴的密度、盖度和空间分布特征, 并分析了土壤理化环境与蚂蚁种类总数、巢穴密度及盖度的相关性。结果表明, 不同演替阶段热带森林蚂蚁种类总数、巢穴的密度及盖度大小顺序为: 小果野芭蕉 (Musa acuminata)群落>白背桐(Mallotus paniculatus)群落>思茅崖豆(Mellettia leptobotrya)群落, 并且热带森林的演替类型显著影响蚂蚁种类总数及巢穴密度, 而对巢穴盖度的影响未达到显著水平; 蚂蚁种类总数、巢穴密度与土壤总有机碳和水解氮显著正相关, 与土壤容重和土壤含水率显著负相关, 但所选择的土壤理化指标与巢穴盖度的相关性均未达到显著水平; 蚂蚁巢穴的空间分布呈随机分布格局。我们的数据表明, 不同演替阶段热带森林所形成的植被类型及土壤环境状况共同影响定居的蚂蚁种类总数与筑巢密度。     

Functional and phylogenetic approaches reveal the evolution of diversity in a hyper diverse biota

Ant communities in tropical forests may be governed by varying assembly mechanisms, depending on the particular habitat investigated. We compared phylogenetic diversity and structure across two forest biomes (dry and humid) and two vertical layers (arboreal and terricolous) in ant communities in Madagascar, and assessed the influence of invasive species on this community structure. We estimated phylogenetic signal and correlated evolution for habitat and several functional traits and tested for conservatism in relevant functional and habitat traits. Ancestral states were reconstructed to illuminate the evolution of habitat traits. All analyses utilized phylogenies estimated from newly generated data from three nuclear markers for 290 Malagasy ant taxa. Dry forests, although lower in species richness, were found to support equally high lineage diversity as humid forests. In contrast, phylogenetic diversity was much lower in arboreal than in terricolous communities. We observed significant phylogenetic clustering in the combined humid forest and in the arboreal–humid, arboreal–dry and terricolous–humid communities, whereas the combined dry forest community was overdispersed. Among ant communities in Madagascar, overdispersion and competition therefore may be more prevalent in dry forest, and habitat filtering may be more dominant in humid forest. Excluding invasive ant species had little overall effect on community structure. All investigated traits showed low to intermediate conservatism; strong support for correlated evolution was found for increased eye size and an arboreal lifestyle. Habitat transitions from humid to dry and from terricolous to arboreal occurred more frequently, and ancestors of most lineages were predicted to be terricolous or humid‐forest adapted. We conclude that most Malagasy ant clades first colonized humid forests and subsequently transitioned into dry forests, indicating that previous hypotheses on the evolution of Madagascar's hyperdiverse biota may not apply to ants and other arthropods.     

Effects of canopy connectivity on the arboreal ant community in coffee shade trees

Canopy connectivity influences foraging, movement, and competition in arboreal ant communities. Understanding how canopy connectivity affects arboreal ant communities could inform the development of management practices that maximize services from known biocontrol agents. We experimentally manipulated connectivity between the crowns of large shade trees to investigate the effects of canopy connectivity on arboreal ant species richness and composition in a coffee agroecosystem. A linear mixed-effects analysis showed that the number of species observed at baits set in tree crowns increased significantly after the crowns had been connected with nylon ropes. Crowns that were connected increased in similarity of ant species composition, particularly between adjacent connected crowns. Connectivity may increase the number of species present in tree crowns by allowing ants to move and forage in the canopy while bypassing trunks with more aggressive, territorial species such as Azteca sericeasur. Because twig-nesting species in the upper canopy have been shown to act as biocontrol agents of herbivores, an increase in species richness in tree crowns could have positive implications for agricultural pest-control services.     

Spatial Variation in Army Ant Swarm Raiding and its Potential Effect on Biodiversity

Invertebrate communities of the tropical rain forest floor are highly diverse, characterized by patchy species distribution patterns and high variation in species density. Spatial variation in the foraging activity of swarm raiding army ants, prime invertebrate predators in tropical rain forests, is discussed as a mechanism contributing to these patterns, but highly resolved long‐term data on army ant raiding on the local and landscape scale are hitherto lacking. In this study, 196 positions in 11 study sites in a tropical rain forest in western Kenya were continuously monitored over ~4 mo for the occurrence of swarm raids of army ants. Using population simulation analyses, the consequences of army ant raiding for prey communities were assessed. We found an unexpectedly high variation in raid rates at the study site and landscape scale. The weekly chance of communities to become raided by army ants was on average 0.11, but ranged from 0 to 0.50 among the 196 positions. Simulating population developments of two Lotka–Volterra species—showing slight trade‐offs between competitive strength and resistance to army ant raids—in the real raiding landscapes showed that the observed spatial variation in raid rates may produce high prey diversity at larger spatial scales (due to high β‐diversity) and strong variation in species density. Our results indicate that high spatial variation in army ant swarm raiding is a mechanism capable of generating patchy species distribution patterns and maintaining the high biodiversity of invertebrate communities of the tropical rain forest floor.     

The contribution of cacao agroforests to the conservation of lower canopy ant and beetle diversity in Indonesia

The ongoing destruction of tropical rainforests has increased the interest in the potential value of tropical agroforests for the conservation of biodiversity. Traditional, shaded agroforests may support high levels of biodiversity, for some groups even approaching that of undisturbed tropical forests. However, it is unclear to what extent forest fauna is represented in this diversity and how management affects forest fauna in agroforests. We studied lower canopy ant and beetle fauna in cacao agroforests and forests in Central Sulawesi, Indonesia, a region dominated by cacao agroforestry. We compared ant and beetle species richness and composition in forests and cacao agroforests and studied the impact of two aspects of management intensification (the decrease in shade tree diversity and in shade canopy cover) on ant and beetle diversity. The agroforests had three types of shade that represented a decrease in tree diversity (high, intermediate and low diversity). Species richness of ants and beetles in the canopies of the cacao trees was similar to that found in lower canopy forest trees. However, the composition of ant and beetle communities differed greatly between the agroforest and forest sites. Forest beetles suffered profoundly from the conversion to agroforests: only 12.5% of the beetle species recorded in the forest sites were also found in the agroforests and those species made up only 5% of all beetles collected from cacao. In contrast, forest ants were well represented in agroforests, with 75% of all species encountered in the forest sites also occurring on cacao. The reduction of shade tree diversity had no negative effect on ants and beetles on cacao trees. Beetle abundances and non-forest ant species richness even increased with decreasing shade tree diversity. Thinning of the shade canopy was related to a decrease in richness of forest ant species on cacao trees but not of beetles. The contrasting responses of ants and beetles to shade tree management emphasize that conservation plans that focus on one taxonomic group may not work for others. Overall ant and beetle diversity can remain high in shaded agroforests but the conservation of forest ants and beetles in particular depends primarily on the protection of natural forests, which for forest ants can be complemented by the conservation of adjacent shaded cacao agroforests.     

Effects of an Epiphytic Orchid on Arboreal Ant Community Structure in Panama

Epiphytes are conspicuous structural elements of tropical forest canopies. Individual tree crowns in lowland forests may support more than 30 ant species, yet we know little about the effects of epiphytes on ant diversity. We examined the composition of arboreal ant communities on Annona glabra trees and their interactions with the epiphytic orchid Caularthron bilamellatum in Panama. We surveyed the ants on 73 trees (45 with C. bilamellatum and 28 lacking epiphytes) and recorded their nest sites and behavioral dominance at baits. We found a total of 49 ant species (in 20 genera), ranging 1–9 species per tree. Trees with C. bilamellatum had higher average (±SD) ant species richness (4.2±2.28) than trees without epiphytes (2.7±1.21). Hollow pseudobulbs (PBs) of C. bilamellatum were used as nest sites by 32 ant species, but only 43 percent of suitable PBs were occupied. Ant species richness increased with PB abundance in trees, but nest sites did not appear to be a limiting resource on A. glabra. We detected no close association between ants and the orchid. We conclude that higher ant species richness in the presence of the orchid is due to bottom‐up effects, especially the year‐round supply of extrafloral nectar. The structure of ant communities on A. glabra partly reflects interference competition among behaviorally dominant species and stochastic factors, as observed in other forests.     

Numerically dominant species drive patterns in resource use along a vertical gradient in tropical ant assemblages

Resource availability can influence the foraging strategy adopted by different ant species as they endeavor to meet nutrient demands of the colony. In tropical rain forests, environmental conditions including resource availability vary over a vertical gradient. Consequently, nitrogen is predicted to become more limiting than carbohydrates toward the canopy as food webs shift to become more reliant on plant-based resources. We used a “bait-choice” experiment in a tropical rain forest to examine differences in protein and carbohydrate use with height and determined whether there were differences in response between common (numerically dominant) and rare species. Additionally, we investigated the nutrient use at the species level. Using species co-occurrence analysis, we examined interspecific competition by testing the co-occurrence of ant species at the tree level. Over the 12 trees investigated, 124 morphospecies were identified with eight species comprising 90% of total ant abundance. Species richness and protein use increased with height of bait for all species pooled and for common species but not rare species. Correspondingly, relative carbohydrate use decreased with height. We found greater species richness of rare species on carbohydrate baits compared with protein baits. Ant species were randomly distributed among trees when all species were included in co-occurrence analysis. However, when only common species were considered, segregation between species was evident among trees providing evidence for the presence of ant mosaics. Our results suggest that nitrogen limitation in the canopy may not be true for the whole ant assemblage but rather for the few common species.     

Convergence in community structure and dietary adaptation in Australian possums and gliders and Malagasy lemurs

Abstract The possums and gliders of Australia and the lemurs of Madagascar are ideally suited for investigation of radiation and convergence in arboreal mammal communities because of their long history of independent radiation in similar biophysical environments. Possum and glider communities at 22 sites in Australia and lemur communities at 18 sites in Madagascar were compared to evaluate patterns of community and species convergence in two shared habitats (evergreen and deciduous monsoon rainforests) and three derived habitats (Australian eucalypt forests and heaths, and Malagasy thorn scrubs). Arboreal mammals were classified into five dietary guilds for comparison of community structure and species adaptation. There is little evidence of convergence in trophic structure at the community level, but strong evidence of convergence in dietary adaptation at the species and niche level. Malagasy rainforest communities were characterized by a higher proportion of frugivore-folivores, and Australian rainforest and moist eucalypt communities by a higher proportion of folivores and folivore-frugivores. Possum and glider communities in temperate eucalypt forests and heaths supported a higher proportion of nectarivores and exudivores. Overall species richness was significantly higher in Madagascar. Four hypotheses are erected to account for these differences. The relative scarcity of frugivores in Australian evergreen rainforests is accounted for by the predominance of bird-dispersed fruits, and fruiting phenologies unsuitable for mammal exploitation and dispersal. Higher folivore diversity in Australian evergreen rainforests is associated with a more pronounced seasonal shortage of edible fruits and new leaf, which has increased dependence on mature leaf. Folivore species richness is greatest in Australian moist eucalypt forests where structural complexity, sustained by serai responses to wildfire, permits a high level of vertical segregation. Increased nectarivory and exudivory in Australian temperate eucalypt forests and heaths is associated with Mediterranean winter rainfall regimes, which permit year round exudate production, and not with the absence of nectar feeding bats as previously supposed. A lower overall species richness in Australian rainforests is attributed to a longer history of contraction and fragmentation by anthropogenic fires (monsoon rainforests) and Pleistocene climatic change (tropical evergreen rainforests). A high degree of convergence is apparent between genera occupying folivore and wood gouging niches, in terms of gastrointestinal morphology (e.g. Pseudocheirus-Lepilemur) and dental morphology (e.g. Dactylopsila-Daubentonia). Divergences are most apparent in adaptations associated with frugivory, including larger body size, diurnality, and bipedal suspension and leaping modes of locomotion (in Eulemur spp.). This study highlights the importance of founding effects, competition from other vertebrate taxa, coevolution between animals and their food plants, and differences in biophysical environments, as determinants of mammalian radiation and convergence.     

Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

Optimizing Diversity Assessment Protocols for High Canopy Ants in Tropical Rain Forest

The canopies of tropical rain forests support highly diverse, yet poorly known, animal and plant communities. It is vital that researchers who invest the time needed to gain access to the high canopy are able efficiently to survey the animals and plants that they find there. Here, we develop diversity assessment protocols for one of the most ecologically important canopy animal groups, the ants, in lowland dipterocarp rain forest in Sabah, Malaysia. We design and test a novel trap (the purse‐string trap) that can be remotely collected, thus avoiding disturbance to ants. We compare this modified trap with two other methods for surveying canopy ants: precision insecticide fogging and baited pitfall trapping. In total, we collected 39,351 ants belonging to 173 species in 38 genera. Fogging collected the most individuals and species, followed by purse‐string trapping with baited pitfall trapping catching the fewest. Fogging also resulted in samples with a different species composition to purse‐string trapping and baited pitfall trapping, which were not different from one another. Using a ‘greedy algorithm’, which guides the selection of inventory methods in order to maximize new species discovered per researcher‐hour, we show that projects allocating fewer than 132 researcher‐hours to canopy ant collection and identification should sample exclusively using fogging. Those with more time should use a combination of methods. This prioritization technique could be used to accelerate species discovery in future rapid biodiversity assessments. Abstract in Malay is available in the online version of this article.     

Arboreal ants as key predators in tropical lowland rainforest trees

Ants numerically dominate the canopy fauna of tropical lowland rain forests. They are considered to be key predators but their effects in this regard have only rarely been studied on non-myrmecophytes. A conspicuously low abundance of less mobile, mainly holometabolous arthropods like Lepidoptera larvae corresponds with ant dominance, while hemimetabolous highly mobile nymphs occur regularly and in large numbers in the trees. This is in contrast to the temperate regions where ants are mostly lacking on trees and holometabolous larvae are frequent. In this study we experimentally measured ant predation in the trees by offering caterpillars as baits. Fifty-four ant species were tested, of which 46 killed caterpillars and carried them away to their nests while only eight species ignored the offered larvae. Insecticidal knockdown fogging of ten trees after finishing the prey experiments showed that on average 85% of ant individuals per tree were predacious. With the analysis of another 69 foggings and meticulous observations in many other trees this suggests that arboreal ants are responsible for the low abundance of less mobile arthropods in tropical lowland rain forest canopies. Ant predation was significantly lower in a disturbed forest indicating that human disturbance induces a change in the functional interactions in these ecosystems.