Arboreal Ants Research Articles

Arboreal Ant Species Richness in Primary Forest, Secondary Forest, and Pasture Habitats of a Tropical Montane Landscape1

ABSTRACTCanopy invertebrates may reflect changes in tree structure and microhabitat that are brought about by human activities. We used the canopy fogging method to collect ants from tree crowns in primary forest, secondary forest, and pasture in a Neotropical cloud forest landscape. The total number of species collected was similar in primary forest (21) and pasture (20) habitats, but lower in secondary forest (9). Lower diversity in secondary forest was caused by lower species density (no. of species per sample). Rarefaction curves based on number of species occurrences suggest similar community species richness among the three habitats. This study has implications for conservation of tropical montane habitats in two ways. First, arboreal ant species density is reduced if secondary forest replaces primary forest, which increases the chance of extinction among rare species. Second, pasture trees may serve as repositories of primary forest ant communities due to similar tree structure.RESUMENLos disturbios antropogénicos sobre la estructura de los árboles y microhabitat afectara la diversidad de invertebrados del dosel. Se usó la técnica de fumigación para colectar hormigas de las copas de los árboles en tres ambientes en un bosque nuboso Neotropical; bosque primario, bosque secundario, y pastizal. La cantidad total de especies colectadas fue similar entre el bosque primario (21) y el pastizal (20), pero menor en el bosque secundario (9). La baja riqueza en el bosque secundario fue consequencia de menor densidad de especies (cantidad de especies por muestra). Las curvas de rarefacción basada en presencia/ausencia de especies sugieren que la riqueza a nivel de la comunidad es similar entre los tres tipos de hábitat. Estos resultados puede aportar a la conservación de los hábitat tropicales nubosos de dos maneras. Primero, sugieren que la densidad de especies de hormigas es baja cuando un bosque primario es reemplazado por un bosque secundario. Segundo, los árboles grandes de pastizales pueden servir como refugios para las comunidades de hormigas, debido a que presentan similitud con la estructura de un bosque primario.

Arboreal Ant Species Richness in Primary Forest, Secondary Forest, and Pasture Habitats of a Tropical Montane Landscape1

Canopy invertebrates may reflect changes in tree structure and microhabitat that are brought about by human activities. We used the canopy fogging method to collect ants from tree crowns in primary forest, secondary forest, and pasture in a Neotropical cloud forest landscape. The total number of species collected was similar in primary forest (21) and pasture (20) habitats, but lower in secondary forest (9). Lower diversity in secondary forest was caused by lower species density (no. of species per sample). Rarefaction curves based on number of species occurrences suggest similar community species richness among the three habitats. This study has implications for conservation of tropical montane habitats in two ways. First, arboreal ant species density is reduced if secondary forest replaces primary forest, which increases the chance of extinction among rare species. Second, pasture trees may serve as repositories of primary forest ant communities due to similar tree structure.

Tree heterogeneity, resource availability, and larger scale processes regulating arboreal ant species richness

Abstract Processes acting on different spatial and temporal scales may influence local species richness. Ant communities are usually described as interactive and therefore determined by local processes. In this paper we tested two hypotheses linked to the question of why there is local variation in arboreal ant species richness in the Brazilian savanna (‘cerrado’). The hypotheses are: (i) there is a positive relationship between ant species richness and tree species richness, used as a surrogate of heterogeneity; and (ii) there is a positive relationship between ant species richness and tree density, used as a surrogate of resource availability. Arboreal ants were sampled in two cerrado sites in Brazil using baited pitfall traps and manual sampling, in quadrats of 20 m × 50 m. Ant species richness in each quadrat was used as the response variable in regression tests, using tree species richness and tree density as explanatory variables. Ant species richness responded positively to tree species richness and density. Sampling site also influenced ant species richness, and the relationship between tree density and tree species richness was also positive and significant. Tree species richness may have influenced ant species richness through three processes: (i) increasing the variety of resources and allowing the existence of a higher number of specialist species; (ii) increasing the amount of resources to generalist species; and (iii) some other unmeasured factor may have influenced both ant and tree species richness. Tree density may also have influenced ant species richness through three processes: (i) increasing the amount of resources and allowing a higher ant species richness; (ii) changing habitat conditions and dominance hierarchies in ant communities; and (iii) increasing the area and causing a species–area pattern. Processes acting on larger scales, such as disturbance, altitude and evolutionary histories, as well as sampling effect may have caused the difference between sites.

Influence of insecticide treatments on ant-hemiptera associations in tropical plantations.

In this survey conducted in southern Cameroon, we compared ant-Hemiptera associations on plantations treated with insecticides, on plantations 2 years after insecticide treatments ceased, and on control lots that never received insecticide treatments. By eliminating arboreal-nesting ants, insecticides favored the presence of "ecologically dominant" ground-nesting, arboreal-foraging species that occupied the tree crowns. The reinstallation of arboreal ants was slow as 2 yr after insecticide treatment ceased differences with the control lots were significant. This intermediary period also illustrated that arboreal ants can found and develop colonies on trees occupied by ground-nesting species. Certain arboreal species were more frequent during this intermediary period than on the control lots, showing that the period of installation in the trees was followed by competition between arboreal ants. We confirm that ground-nesting ants tend a wide range of hemipteran families, including well known agricultural pests, whereas arboreal ants, particularly dominant species, were mostly associated with Coccidae and Stictococcidae that do not pose problems to the supporting trees. A tree effect was also noted for both ant and hemipteran distribution. We concluded that because of insecticide use, ground-nesting ants pose problems through their associated Hemiptera. On the contrary, dominant arboreal ants, strong predators, benefit their supporting trees by excluding ground-nesting species and tending mostly nonpest Hemiptera. Nevertheless, certain of them, carpenter species or species likely to tend Pseudococcidae, have to be eliminated through integrated management.

Influence of Insecticide Treatments on Ant-Hemiptera Associations in Tropical Plantations

Abstract In this survey conducted in southern Cameroon, we compared ant-Hemiptera associations on plantations treated with insecticides, on plantations 2 years after insecticide treatments ceased, and on control lots that never received insecticide treatments. By eliminating arboreal-nesting ants, insecticides favored the presence of “ecologically dominant” ground-nesting, arboreal-foraging species that occupied the tree crowns. The reinstallation of arboreal ants was slow as 2 yr after insecticide treatment ceased differences with the control lots were significant. This intermediary period also illustrated that arboreal ants can found and develop colonies on trees occupied by ground-nesting species. Certain arboreal species were more frequent during this intermediary period than on the control lots, showing that the period of installation in the trees was followed by competition between arboreal ants. We confirm that ground-nesting ants tend a wide range of hemipteran families, including well known agric...

Do non-myrmocophilic epiphytes influence community structure of arboreal ants?

Abstract In a one-year-survey in Panama we examined the influence of a tree crown's epiphyte assemblage on its ant fauna. Ants were collected with various types of insect traps in 25 crowns of Annona glabra trees. The study trees were assigned to three different categories according to their epiphyte load, and to an epiphyte-free control group. We collected 22,335 specimens of 91 morphospecies, 32 genera and six subfamilies. By far the most abundant species was Solenopsis zeteki, a minute Myrmicinae, which was found in each of the 25 study trees. Many other species were also rather common and widely distributed throughout the study area. Only six species were singletons. Measures of α- and β-diversity, species abundance and species composition were not affected by the epiphyte load of a tree. We also made direct in situ observations of ants on 34 additional Annona glabra trees with and without epiphytes. Workers were attracted with tuna and sugar baits, and interspecific interactions and nesting sites were recorded. In total, 40 species of ants were found, all of which had also been collected in the traps. Almost half of the colonies (48%) used dead wood as nesting substrate, while 29% nested in epiphytes. Consistent with the results of the trap survey, the epiphyte load of the study trees had no influence on ant species richness and composition, but a significant correlation between ant abundance and epiphyte load was detectable. In both data sets, the lack of associations between ant species indicated that the ant assemblages were not structured in a mosaic-like fashion. We conclude that epiphytes do hardly influence the composition of ant assemblages in the studied tree crowns, probably because arboreal ants are highly opportunistic with respect to their host plants. In einer einjahrigen Studie wurde der Einfluss von Epiphytengemeinschaften in Baumkronen auf die dort lebende Ameisenfauna untersucht. Die Ameisenfauna wurde mit zwei unterschiedlichen Methoden erfasst. Zum einen wurden mit verschiedenen Arthropodenfallen Annona glabra Baume beprobt, die gemas ihres Epiphytenbewuchses in drei verschiedene Kategorien und eine unbewachsene Kontrollgruppe eingeteilt wurden. Insgesamt wurden 22.335 Individuen aus 91 Arten gefangen, die 32 Gattungen und sechs Unterfamilien zuzuordnen waren. Die haufigste Art, Solenopsis zeteki, eine winzige Myrmicinae, wurde auf allen 25 beprobten Baumen gefunden. Viele andere Ameisenarten waren ebenfalls recht weit und gleichmasig uber das Untersuchungsgebiet verbreitet. Nur sechs Arten wurden in Einzelexemplaren gefunden. Weder Abundanz der Arten noch Artenzusammensetzung wurden durch die Art des Epiphytenbewuchses beeinflusst. Zweitens wurden auch in-situ-Beobachtungen an einer zusatzlichen Gruppe von 34 Baumen derselben Baumart durchgefuhrt. Diese Baume waren entweder mit Epiphyten bewachsen oder epiphytenfrei. Arbeiterinnen wurden mit Thunfisch- und Zuckerkodern angelockt, und zwischenartliche Interaktionen sowie die Neststandorte registriert. In dieser Teiluntersuchung wurden 40 Arten bestimmt, die ausnahmslos auch in den Fallen gefunden worden waren. Etwa die Halfte der Kolonien (48%) befand sich in Totholz und 29% in Epiphyten. Auch hier zeigte sich kein Einfluss des Epiphytenbewuchses auf die Zahl der Arten oder ihre Zusammensetzung, die Korrelation zwischen Ameisenabundanz und Epiphytenbewuchs war aber signifikant. In beiden Teilstudien gab es keine signifikanten Arten-Assoziationen, die Ameisengemeinschaften waren also nicht mosaikartig strukturiert. Aus den Daten kann die Schlussfolgerung abgeleitet werden, dass Epiphyten die Zusammensetzung der Ameisengemeinschaften in den untersuchten Baumkronen kaum beeinflussen, weil Ameisen wahrscheinlich sehr opportunistisch auf unterschiedliche pflanzliche Ressourcen reagieren.

Seasonal Foraging Activity and Bait Preferences of Ants on Barro Colorado Island, Panama1

ABSTRACTA yearlong arboreal baiting survey of ants was conducted during 1983 on Barro Colorado Island, Panama. Because of a severe El Nino event, the 1983 dry season in Panama was exceptionally long and dry with a distinct boundary between the dry and wet seasons. Baits, located on tree trunks, attracted both terrestrial and arboreal ants, allowing comparisons between the two groups. Species composition at baits changed dramatically with season. Baits were primarily occupied by arboreal species during the dry season, while wet season baits were occupied mostly by terrestrial species. Arboreal and terrestrial ants differed markedly in their preferences for protein‐ or carbohydrate‐based baits; arboreal ants preferred protein‐based baits and terrestrial ants preferred carbohydrate‐based baits. Foraging preference for protein suggests that protein resources were limiting for arboreal ants, particularly during the dry season, and that carbohydrate resources were limiting for terrestrial ants. Fundamental differences in arboreal and terrestrial habitats may promote the differences in foraging strategies observed during an annual cycle in a seasonal tropical forest.

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.

Seasonal Foraging Activity and Bait Preferences of Ants on Barro Colorado Island, Panama1

A yearlong arboreal baiting survey of ants was conducted during 1983 on Barro Colorado Island, Panama. Because of a severe El Niño event, the 1983 dry season in Panama was exceptionally long and dry with a distinct boundary between the dry and wet seasons. Baits, located on tree trunks, attracted both terrestrial and arboreal ants, allowing comparisons between the two groups. Species composition at baits changed dramatically with season. Baits were primarily occupied by arboreal species during the dry season, while wet season baits were occupied mostly by terrestrial species. Arboreal and terrestrial ants differed markedly in their preferences for protein- or carbohydrate-based baits; arboreal ants preferred protein-based baits and terrestrial ants preferred carbohydrate-based baits. Foraging preference for protein suggests that protein resources were limiting for arboreal ants, particularly during the dry season, and that carbohydrate resources were limiting for terrestrial ants. Fundamental differences in arboreal and terrestrial habitats may promote the differences in foraging strategies observed during an annual cycle in a seasonal tropical forest.

North—South Patterns within Arboreal Ant Assemblages from Rain Forests in Eastern Australia1

ABSTRACTThis paper describes the ant assemblages sampled from rain forest canopies ranging from southern Victoria through to Cape York Peninsula, Australia, and also in Brunei. Specifically, it examines the influence of decreasing latitude and variations in elevation on the character, richness, and abundance of the arboreal rain forest ant fauna, and also the relative contribution of ants to the total arthropod community. The sites that were examined included: cool temperate Nothofagus cunninghamii forest from a range of locations in Victoria; cool temperate N. moorei forest at both Werrikimbe and Styx River, New South Wales; notophyll vine forest in Lamington National Park, southeast Queensland; high elevation notophyll vine forest in Eungella National Park, central Queensland; complex notophyll vine forest at Robson Creek, Atherton Tablelands, north Queensland; complex mesophyll vine forest at Cape Tribulation, north Queensland; and mixed dipterocarp forest in Brunei. Although these sites represent a gradient increasingly tropical in character, botanically speaking, Eungella is less tropical than Lamington because of its high elevation. All samples were obtained by fogging the canopy with a rapid‐knockdown pyrethrin pesticide. In all cases, circular funnels were suspended beneath the foliage of individual trees or small plots of mixed canopy. Arthropods were collected four hours after fogging. Following ordinal sorting, ants were identified and counted to morphospecies level. The resulting catch were then standardized across sites as numbers caught per 0.5 m2 sampling funnel. Generic and species richness were higher at the lowland tropical Cape Tribulation sites than at the sites to the south and was comparable with values in the Brunei site. Species richness was negatively correlated with latitude and elevation. Within the Australian rain forest, the lowland/highland break appears to be the strongest predictor of ant relative abundance, with a weaker latitudinal relationship superimposed.

North–South Patterns within Arboreal Ant Assemblages from Rain Forests in Eastern Australia1

This paper describes the ant assemblages sampled from rain forest canopies ranging from southern Victoria through to Cape York Peninsula, Australia, and also in Brunei. Specifically, it examines the influence of decreasing latitude and variations in elevation on the character, richness, and abundance of the arboreal rain forest ant fauna, and also the relative contribution of ants to the total arthropod community. The sites that were examined included: cool temperate Nothofagus cunninghamii forest from a range of locations in Victoria; cool temperate N. moorei forest at both Werrikimbe and Styx River, New South Wales; notophyll vine forest in Lamington National Park, southeast Queensland; high elevation notophyll vine forest in Eungella National Park, central Queensland; complex notophyll vine forest at Robson Creek, Atherton Tablelands, north Queensland; complex mesophyll vine forest at Cape Tribulation, north Queensland; and mixed dipterocarp forest in Brunei. Although these sites represent a gradient increasingly tropical in character, botanically speaking, Eungella is less tropical than Lamington because of its high elevation. All samples were obtained by fogging the canopy with a rapid-knockdown pyrethrin pesticide. In all cases, circular funnels were suspended beneath the foliage of individual trees or small plots of mixed canopy. Arthropods were collected four hours after fogging. Following ordinal sorting, ants were identified and counted to morphospecies level. The resulting catch were then standardized across sites as numbers caught per 0.5 m2 sampling funnel. Generic and species richness were higher at the lowland tropical Cape Tribulation sites than at the sites to the south and was comparable with values in the Brunei site. Species richness was negatively correlated with latitude and elevation. Within the Australian rain forest, the lowland/highland break appears to be the strongest predictor of ant relative abundance, with a weaker latitudinal relationship superimposed.

Food habits of Aenictus army ants and their effects on the ant community in a rain forest of Borneo

In a rain forest at the foot of Mt. Kinabalu, Borneo, we observed the food habits of Aenictus laeviceps and A. gracilis and surveyed the effects of A. laeviceps on the general ant community. We collected totals of 2,360 and 6,268 prey units, respectively, from seven A. laeviceps and six A. gracilis colonies; ants constituted 99.9% of all prey units. Dominant prey genera were Camponotus (48.2% in wet weight), Pseudolasius (20.8%), and Polyrhachis (15.2%) in A. laeviceps and Technomymex (52.1%), Paratrechina (22.4%), and Crematogaster (11.9%) in A. gracilis. A. laeviceps primarily hunted larger ground ants with small- or medium-sized colonies; in contrast, A. gracilis frequently foraged on smaller arboreal ants with larger colonies. Of the total wet weight of prey ants, brood constituted 88.7% in A. laeviceps and only 68.7% in A. gracilis. This was probably because workers of A. gracilis were quicker in behavior and hunted adult workers more frequently than A. laeviceps. While 98.5% of the prey units were transported by single workers of A. gracilis, 71.5% were carried by up to 20 A. laeviceps workers. The numbers of ant colonies and species were significantly smaller in laeviceps-raided quadrats than in controls. This was mostly due to the escape of ants that nested under litter and had relatively larger colonies. Large colonies usually influence the foundation and establishment of incipient small colonies.

Sterilization and canopy modification of a swollen thorn acacia tree by a plant-ant

Obligate symbioses between specialized arboreal ants and plants have evolved independently in many lineages1,2. Ant-plants (myrmecophytes) typically provide hollow nest cavities and nutrition to the occupying ant colony1,3,4,5,6. In turn, resident plant-ants often protect their hosts from herbivory7,8,9,10,11 and/or overgrowth by surrounding vegetation12,13. As individual plants are rarely occupied by more than one ant colony14,15,16,17, co-occurring plant-ant species compete intensely for hosts13,14,18,19. In such multi-species systems, ecological interactions among potential partners may lead to the evolution of cheating20,21. Previous studies have revealed that some specialized plant-ants are effectively parasites of their host-plants8,18,22,23, but the selection pressures favouring such behaviours are poorly understood. Here we describe host parasitism in an east African plant-ant that prunes and sterilizes its host-tree canopies, apparently to minimize contact with competitively dominant ants occupying neighbouring trees. We propose that the high density of ant-trees and low diversity of tree species in this savanna habitat have selected for induced, parasitic pruning of host trees by this competitively subordinate ant species.

Systematics, biogeography and host plant associations of the Pseudomyrmex viduus group (Hymenoptera: Formicidae), Triplaris - and Tachigali -inhabiting ants

The Pseudomyrmex viduus group is a Neotropical clade of arboreal ants containing 13 species, of which three are newly described here P. insuavis, P. ultrix, and P. vitabilis. Most species inhabit the domatia of specialized ant-plants. The ants keep brood and scale insects (Coccoidea) in the plant cavities, and defend their nest sites much more aggressively than do generalist species of Pseudomyrmex nesting in dead twigs. Five species are obligate associates of trees in the genus Triplaris (Polygonaceae) and five taxa are restricted to Tachigali (Fabaceae: Caesalpinioideae). One species, P. viduus (F. Smith), is much less host-specific, inhabiting Cordia, Coussapoa, Ocotea, Pseudobombax, Pterocarpus, Sapium, Triplaris and other myrmecophytes. Nothing is known about the biology of P. vitabilis, a close relative of P. viduus. The last member of the species group, P. kuenckeli (Emery), is a non-specialist but aggressive species that nests in dead branches of various plants. A cladistic analysis indicates that this aggressive behaviour evolved before obligate associations with specialized ant-plants, and that the Triplaris andTachigali inhabitants each form their own clade. P. viduus, which is nested within the Triplaris -associated clade, suggests a possible model for host plant evolution in these ants wherein shifts from one ant-plant to another involve an intermediate phase of expanded host plant use. At least nine other Pseudomyrmex species, from two different species groups (not closely related to the P. viduus group), have evolved specialized associations with Triplaris or Tachigali including five new species P. crudelis, P. deminutus, P. eculeus, P. ferox and P. hospitalis. Although the P. viduus group is centred in the Amazon basin, the geographic ranges of most species do not coincide with the Pleistocene forest refugia proposed by Haffer and others. A consideration of the phylogenetic relationships, distribution patterns, and host plant specificity of the ants indicates that much of the diversification of the P. viduus group occurred before the Pleistocene, and that the interactions with Triplaris and Tachigali plants are also of Tertiary origin.

Systematics, biogeography and host plant associations of the Pseudomyrmex viduus group (Hymenoptera: Formicidae), Triplaris- and Tachigali-inhabiting ants

The Pseudomyrmex viduus group is a Neotropical clade of arboreal ants containing 13 species, of which three are newly described here: P. insuavis, P. ultrix, and P. vitabilis. Most species inhabit the domatia of specialized ant-plants. The ants keep brood and scale insects (Coccoidea) in the plant cavities, and defend their nest sites much more aggressively than do generalist species of Pseudomyrmex nesting in dead twigs. Five species are obligate associates of trees in the genus Triplaris (Polygonaceae) and five taxa are restricted to Tachigali (Fabaceae: Caesalpinioideae). One species, P. viduus (F. Smith), is much less host-specific, inhabiting Cordia, Coussapoa, Ocotea, Pseudobombax, Pterocarpus, Sapium, Triplaris and other myrmecophytes. Nothing is known about the biology of P. vitabilis, a close relative of P. viduus. The last member of the species group, P. kuenckeli (Emery), is a non-specialist but aggressive species that nests in dead branches of various plants. A cladistic analysis indicates that this aggressive behaviour evolved before obligate associations with specialized ant-plants, and that the Triplaris and Tachigali inhabitants each form their own clade. P. viduus, which is nested within the Triplarisassociated clade, suggests a possible model for host plant evolution in these ants wherein shifts from one ant-plant to another involve an intermediate phase of expanded host plant use. At least nine other Pseudomyrmex species, from two diVerent species groups (not closely related to the P. viduus group), have evolved specialized associations with Triplaris or Tachigali including five new species: P. crudelis, P. deminutus, P. eculeus, P. ferox and P. hospitalis. Although the P. viduus group is centred in the Amazon basin, the geographic ranges of most species do not coincide with the Pleistocene forest refugia proposed by HaVer and others. A consideration of the phylogenetic relationships, distribution patterns, and host plant specificity of the ants indicates that much of the diversification of the P. viduus group occurred before the Pleistocene, and that the interactions with Triplaris and Tachigali plants are also of Tertiary origin.

Impact of tree isolation on arboreal and ground ant communities in cleared pasture in the Atlantic rain forest region of Bahia, Brazil

Summary. The Atlantic rain forest of south-eastern Brazil has been substantially cleared, resulting in the creation of a fragmented landscape. In addition to the small fragments of forest that remain, the pasture is often scattered with isolated trees. This paper investigates the capacity of these isolated trees to support representatives of the original Atlantic rain forest ant communities and also how these arboreal ants interact with the disturbance-associated ant fauna of the pasture beneath them. Twenty trees in the grassland, representing a range of distances from the forest, and 10 trees within the forest, were selected for sampling. Arboreal ants were sampled by hand collection and chemical knockdown, while the ants on the ground beneath were sampled by pitfall traps and Winkler sacks. Pasture trees supported a moderately high richness of arboreal ant species. The richness of ants on pasture trees appears to be independent of distance from forest, although this might become a significant factor on trees that are more isolated than those studied here. Ant species richness on pasture trees is higher if the trees are large, support a high epiphyte load and are native to the area. Isolated trees within the agricultural matrix therefore play some role in conserving elements of the original forest ant fauna. Since some of the species on pasture trees have been observed to reach dominant or sub-dominant status in nearby forest and cocoa farms, they may play some role in limiting pest outbreaks in the pasture close to the trees. If this is so, there may be a case for retaining an adequate density of trees to enable the influence of arboreal ants to extend over as much pasture as possible.

Innate attraction supplants experience during host plant selection in an obligate plant-ant

In order to assess the relative contributions of innate attraction and environmentally-induced preference to nest plant selection by the obligate plant-ant Tetraponera aethiops, we submitted both mature and callow workers to paired choice tests using the leaves of four plant species, including their natural host plant Barteria fistulosa. Mature workers taken from nature as well as mature workers after 25 days of laboratory breeding (with or without contact with B. fifstulosa leaves) always showed a great preference for shelters made with the leaves of B. fistulosa. Similar results were obtained with callow workers isolated from any plant material during the first 25 days of imaginal (=adult) life or during pre-imaginal development. But those reared in contact with test-plant leaves for the same two periods showed significantly less preference for B. fistulosa than did the controls. As a result, experience gained during the larval and nymphal stages as well as during the first part of adult life (i.e. pre-imaginal as well as early learning) was demonstrated in T. aethiops, but in this situation of species-specific interaction it reinforces innate attraction as both larvae and adults live in contact with B. fistulosa in nature. These results are discussed in comparison with previous data collected on arboreal ants not specifically paired with a plant, for which an environmentally-induced preference can supplant innate attraction.

Territorialité et agressivité intra- et interspécifique dans les mosaïques de fourmis arboricoles

Tropical arboreal ants are distributed in a mosaic pattern in the canopy of forests and tree crop plantations each of them characterised by their status of dominance. One can distinguish ‘dominant’ species, characterised by extremely populous societies and highly developed interspecific as well as intraspecific territorial behaviour. They tolerate on their territory nonterritorial and less populous species classified as ‘non-dominants’. Nonetheless, many species do exist whose status is intermediary. Usually, they behave like non-dominant species but are able, under certain conditions, to defend a territory. They are cited as ‘sub-dominant’. According to the chemical trapping method employed by researchers, the structure of mosaics have most often been studied using an index of dominance, characterised by the number of negative or positive associations between one species and the others. This index only covers the relative presence or absence of the different species on the same trees. It only gives a punctual statement on the structure of the mosaic without any notion of evolution of the mosaic in time. It does not take into account the behavioural intra- and interspecific interactions. Aggressive interactions between species depend on genetic and environmental factors. Many studies have shown that aggressiveness is closely related to a mechanism of interindividual discrimination, permitting an individual to discriminate between nestmates and non-nestmates. This colonial recognition is based on the existence of a ‘colonial odour’ as a result of a blending of ‘individual odours’. Each individual odour is due to cuticular hydrocarbons which play the role of a contact pheromone. The colonial odour also depends on the environmental odour of the nest. Aggressiveness which results from this mechanism of recognition can be expressed through different mechanisms such as territorial behaviour, dominance hierarchy, and ritualised aggressive behaviour. Territorial behaviour is the expression of a strong intraspecific aggressiveness, by which workers of a colony defend an area of their vital domain against neighbouring conspecifics. In arboreal ant mosaics, dominance hierarchy can exist between dominant ants, and should explain the overturning of dominant ants in time. Ritualised behaviours were observed under intra- and interspecific low-aggressiveness conditions and allow to economise the loss of one or several workers during fights whose issue are uncertain. Their systematic study would greatly facilitate understanding of the evolution of arboreal mosaics.

L'adaptation à la vie arboricole chez les fourmis

Arboreal ants constitute one of the largest animal groups in the tropical forest canopy not just in terms of their individual number and overall biomass, but also in terms of diversity. Because the arboreal habitat has particular characteristics which clearly differentiate it from the ground, the ants' ecological success depends on developing adaptations designed to overcome the constraints peculiar to this environment in order to best exploit its resources. Although it is not always possible to identify all of the characteristics common to all arboreal ants species, those species said to be ‘dominant’ share morphological, physiological and behavioural traits which differentiate them from other ants. The most important difference is the type of food sources exploited. These dominant species feed principally on the nectar of the extra-floral nectaries of plants or on the honeydew of Homoptera. These products are extremely rich in sugary substances and low in nitrogen. The exploitation of these food sources has, then, repercussions on the growth and reproduction of ant colonies, given the importance of nitrogen as a limiting factor. Obtaining the necessary amount of nitrogen involves exploiting large volumes of liquid. From a purely morphological point of view, it is interesting to note that the digestive system of most of the dominant ant species has a modified proventriculus and a thin cuticle which means less need for proteins. These modifications permit individual foragers to transport large volumes of liquid by holding them in their crops in order to distribute them to the rest of the colony. The venom of many of these species also differs from that of other ants in that it lacks nitrogen. Dominant species have, thus, developed adaptations related to their feeding habits, lowering the need for nitrogen in individuals and, in this way, permitting the greater development of the colony. The exploitation of this type of food source is also facilitated by the type of nests constructed by the ants. Indeed, the elaboration of a nest independently of any structure provided by the plant offers numerous advantages, despite the important energetic costs involved. These types of nests allow the ant to (1) ‘choose’ the nest site and thus to install the nest on plants providing the best resources (extra-floral nectaries or Homoptera attendance) and (2) exploit a greater number of resources with lower energy costs by placing parts of the colonies near food sources. In addition, even if part of the sugary food substances are destined for the synthesis of molecules necessary to the survival and proper functioning of individuals, there is a large surplus. This surplus might indirectly reinforce these species' defensive systems on both an inter- as well as intraspecific level by serving as an energy source enabling individuals to have high level of activity and aggressiveness. The remaining arboreal species, which — in terms of diversity — make up the majority, do not share common traits which permit them to be clearly differentiated from terricolous species. Nevertheless, comparisons conducted within a genus show that certain arboreal species have developed adaptations to their way of life, and not only those species living in the domatia of ant plants, but also other, strictly arboreal species. As is the case for dominant species, the modifications observed originate especially in response to the two principal limiting factors in the arboreal habitat: available food resources and a limited number of nest sites.

Tropical arboreal ant mosaics: innate attraction and imprinting determine nest site selection in dominant ants

The modalities of nest site selection have, until now, been a key factor missing in the understanding of the arboreal ant mosaic, the manipulation of which could be used to favour one ant species to the detriment of others in biological control. We compared two dominant African arboreal ants of economic importance, Tetramorium aculeatum (Myrmicinae) and Oecophylla longinoda (Formicinae). The two species differed in terms of innate attraction to nesting site plants, their hierarchies of attractiveness being nearly inverse. Winged females and workers were confronted with choice tests using four plant species. By using winged females and workers originating from one of the plants to be tested, we showed the existence of a familiarisation process which can supersede innate attraction in both species. We recorded the same effect in neonatal workers bred in the laboratory in contact with a tested plant for 25 days after emergence, while mature workers could not be conditioned. There is, therefore, early learning, with a sensitive period after which the influence of the environment ceases, suggesting that this is a true imprinting process. Choice tests using neonates produced from larvae and pupae bred in the laboratory in contact with the leaves of each tested plant permitted us to demonstrate the existence of pre-imaginal learning. Nest site selection therefore depends on innate selective attraction and on environmental factors whose effect begins at the larval stage. There is, therefore, the potential to “control” imprinting, allowing one ant species to be favoured to the detriment of others in monospecific tree crop plantations.