Les relations entre fourmis arboricoles et guêpes sociales sous les tropiques

Due to the brood that it contains, the nest of social wasps constitutes a major source of protein for eventual predators. In the tropics, ants are generally considered as the most efficacious among them. In the face of such predation, wasps have developed over the course of evolution a great number of strategies. In addition to guarding and defensive behaviours displayed by workers, social wasps have nests whose structure is a response to the predation pressure exerted by ants. Thus, the presence of a petiole attaching the nest to a natural support itself sometimes selected because of its limited accessibility constitutes a frequent example of architectural defence. The effectiveness of the petiole is often improved through the repeated application of repellent substances on its surface. In the same way, wasps that found their new societies through swarming construct nests protected by an envelope. In the American tropics, army ants which hunt on the ground and can rapidly take over a tree, regularly pillage wasp nests. Because no defensive strategy is efficacious against such predators, the choice of nest site becomes decisive. Paradoxically, it is by nesting in close proximity to an arboreal ant nest that certain wasp species have been able to find a way of responding to the threat exerted by other ants. These ants, which tolerate the presence of the wasp nest, are capable of efficaciously protecting the tree or the branch sheltering the nest.     

An uneasy alliance:a nesting association between aggressive ants and equally fierce social wasps

Although the Neotropical territorially dominant arboreal ant Azteca chartifex Forel is very aggressive towards any intruder,its populous colonies tolerate the close presence of the fierce polistine wasp Polybia rejecta(F.).In French Guiana,83.33%of the 48 P.rejecta nests recorded were found side by side with those of A.chartifex.This nesting association results in mutual protection from predators(i.e.,the wasps protected from army ants;the ants protected from birds).We conducted field studies,laboratorybased behavioral experiments and chemical analyses to elucidate the mechanisms allowing the persistence of this association.Due to differences in the cuticular profiles of the two species,we eliminated the possibility of chemical mimicry.Also,analyses of the carton nests did not reveal traces of marking on the envelopes.Because ant forager flows were not perturbed by extracts from the wasps’Dufour’s and venom glands,we rejected any hypothetical action of repulsive chemicals.Nevertheless,we noted that the wasps"scraped"the surface of the upper part of their nest envelope using their mandibles,likely removing the ants'scent trails,and an experiment showed that ant foragers were perturbed by the removal of their scent trails.This leads us to use the term"erasure hypothesis."Thus,this nesting association persists thanks to a relative tolerance by the ants towards wasp presence and the behavior of the wasps that allows them to"contain"their associated ants through the elimination of their scent trails,direct attacks,"wing-buzzing"behavior and ejecting the ants.     

How to escape from the host nest: Imperfect chemical mimicry in eucharitid parasitoids and exploitation of the ants’ hygienic behavior

Communication in ants is based to a great extent on chemical compounds. Recognition of intruders is primarily based on cuticular hydrocarbon (CHC) profile matching but is prone to being cheated. Eucharitid wasps are specific parasitoids of the brood of ants; the immature stages are either well integrated within the colony or are protected within the host cocoons, whereas adult wasps at emergence must leave their host nest to reproduce and need to circumvent the ant recognition system to escape unscathed. The behavioral interactions between eucharitid wasps and workers of their host, the Neotropical ant Ectatomma tuberculatum, are characterized. In experimental bioassays, newly emerged parasitoids were not violently aggressed. They remained still and were grabbed by ants upon contact and transported outside the nest; host workers were even observed struggling to reject them. Parasitoids were removed from the nest within five minutes, and most were unharmed, although two wasps (out of 30) were killed during the interaction with the ants. We analyzed the CHCs of the ant and its two parasitoids, Dilocantha lachaudii and Isomerala coronata, and found that although wasps shared all of their compounds with the ants, each wasp species had typical blends and hydrocarbon abundance was also species specific. Furthermore, the wasps had relatively few CHCs compared to E. tuberculatum (22–44% of the host components), and these were present in low amounts. Wasps, only partially mimicking the host CHC profile, were immediately recognized as alien and actively removed from the nest by the ants. Hexane-washed wasps were also transported to the refuse piles, but only after being thoroughly inspected and after most of the workers had initially ignored them. Being recognized as intruder may be to the parasitoids’ advantage, allowing them to quickly leave the natal nest, and therefore enhancing the fitness of these very short lived parasitoids. We suggest that eucharitids take advantage of the hygienic behavior of ants to quickly escape from their host nests.     

A Unique Nest-Protection Strategy in a New Species of Spider Wasp

Hymenoptera show a great variation in reproductive potential and nesting behavior, from thousands of eggs in sawflies to just a dozen in nest-provisioning wasps. Reduction in reproductive potential in evolutionary derived Hymenoptera is often facilitated by advanced behavioral mechanisms and nesting strategies. Here we describe a surprising nesting behavior that was previously unknown in the entire animal kingdom: the use of a vestibular cell filled with dead ants in a new spider wasp (Hymenoptera: Pompilidae) species collected with trap nests in South-East China. We scientifically describe the ‘Bone-house Wasp’ as Deuteragenia ossarium sp. nov., named after graveyard bone-houses or ossuaries. We show that D. ossarium nests are less vulnerable to natural enemies than nests of other sympatric trap-nesting wasps, suggesting an effective nest protection strategy, most likely by utilizing chemical cues emanating from the dead ants.     

The Antsy Social Network: Determinants of Nest Structure and Arrangement in Asian Weaver Ants

Asian weaver ants (Oecophylla smaragdina) are arboreal ants that are known to form mutualistic complexes with their host trees. They are eusocial ants that build elaborate nests in the canopy in tropical areas. A colony comprises of multiple nests, usually on multiple trees, and the boundaries of the colony may be difficult to identify. However, they provide the ideal model for studying group living in invertebrates since there are a definite number of nests for a given substrate, the tree. Here, we briefly examine the structure of the nests and the processes involved in the construction and maintenance of these nests. We have described the spatial arrangement of weaver ant nests on trees in two distinct tropical clusters, a few hundred kilometres apart in India. Measurements were made for 13 trees with a total of 71 nests in the two field sites. We have considered a host of biotic and abiotic factors that may be crucial in determining the location of the nesting site by Asian weaver ants. Our results indicate that tree characteristics and architecture followed by leaf features help determine nest location in Asian weaver ants. While environmental factors may not be as influential to nest arrangement, they seem to be important determinants of nest structure. The parameters that may be considered in establishing the nests could be crucial in picking the evolutionary drivers for colonial living in social organisms.     

Arboreal thorn-dwelling ants coexisting on the savannah ant-plant, Vachellia erioloba, use domatia morphology to select nest sites

Nest site selection in arboreal, domatia-dwelling ants, particularly those coexisting on a single host plant, is little understood. To examine this phenomenon we studied the African savannah tree Vachellia erioloba, which hosts ants in swollen-thorn domatia. We found four ant species from different genera (Cataulacus intrudens, Tapinoma subtile, Tetraponera ambigua and an unidentified Crematogaster species). In contrast to other African ant plants, many V. erioloba trees (41 % in our survey) were simultaneously co-occupied by more than one ant species. Our study provides quantitative field data describing: (1) aspects of tree and domatia morphology relevant to supporting a community of mutualist ants, (2) how ant species occupancy varies with domatia morphology and (3) how ant colony size varies with domatia size and species. We found that Crematogaster sp. occupy the largest thorns, followed by C. intrudens, with T. subtile in the smallest thorns. Thorn age, as well as nest entrance hole size correlated closely with ant species occupant. These differing occupancy patterns may help to explain the unusual coexistence of three ant species on individual myrmecophytic trees. In all three common ant species, colony size, as measured by total number of ants, increased with domatia size. Additionally, domatia volume and species identity interact to predict ant numbers, suggesting differing responses between species to increased availability of nesting space. The proportion of total ants in nests that were immatures varied with thorn volume and species, highlighting the importance of domatia morphology in influencing colony structure.     

Strike Fast,Strike Hard: The Red-Throated Caracara Exploits Absconding Behavior of Social Wasps during Nest Predation

Red-throated Caracaras Ibycter americanus (Falconidae) are specialist predators of social wasps in the Neotropics. It had been proposed that these caracaras possess chemical repellents that allow them to take the brood of wasp nests without being attacked by worker wasps. To determine how caracaras exploit nests of social wasps and whether chemical repellents facilitate predation, we: (1) video recorded the birds attacking wasp nests; (2) analyzed surface extracts of the birds'' faces, feet, and feathers for potential chemical repellents; and (3) inflicted mechanical damage on wasp nests to determine the defensive behavior of wasps in response to varying levels of disturbance. During caracara predation events, two species of large-bodied wasps mounted stinging attacks on caracaras, whereas three smaller-bodied wasp species did not. The “hit-and-run” predation tactic of caracaras when they attacked nests of large and aggressive wasps reduced the risk of getting stung. Our data reveal that the predation strategy of caracaras is based on mechanical disturbance of, and damage to, target wasp nests. Caracara attacks and severe experimental disturbance of nests invariably caused wasps to abscond (abandon their nests). Two compounds in caracara foot extracts [sulcatone and iridodial] elicited electrophysiological responses from wasp antennae, and were also present in defensive secretions of sympatric arboreal-nesting Azteca ants. These compounds appear not to be wasp repellents but to be acquired coincidentally by caracaras when they perch on trees inhabited with Azteca ants. We conclude that caracara predation success does not depend on wasp repellents but relies on the absconding response that is typical of swarm-founding polistine wasps. Our study highlights the potential importance of vertebrate predators in the ecology and evolution of social wasps.     

The Role of Wood Ants (Formica rufa group) in Carbon and Nutrient Dynamics of a Boreal Norway Spruce Forest Ecosystem

Wood ants (Formica rufa group) are regarded as keystone species in boreal and mountain forests of Europe and Asia by their effect on ecosystem carbon (C) and nutrient pools and fluxes. To quantify the impact of their activity on boreal forest ecosystems, C, nitrogen (N), phosphorus (P), potassium (K) and calcium (Ca) pools and fluxes in wood ant nests (WAN), and soil were assessed along a 5-, 30-, 60-, and 100-year-old Norway spruce (Picea abies L. Karsten) dominated successional gradient in eastern Finland. Amounts of C and nutrients in WAN increased with stand age, but contained less than 1% of total C and nutrient pools in these stands. The CO₂-efflux from nests was also insignificant, as compared to CO₂-efflux from the forest floor. Annually, the amount of C brought by wood ants into their nests as honeydew, prey and nest-building materials ranged from 2.7 to 49.3 kg ha^?1 C, but this is only 0.1–0.7% of the combined net primary production of trees and understorey in boreal forests. The difference between wood ant nest C inputs and outputs was very small in the younger-aged stands, and increased in the older stands. Carbon accumulation rates in nests over a 100 year period are estimated to be less than 10 kg ha^?1 a^?1. In contrast to C, annual inputs of N, P, and K are larger compared to wood ant nest nutrient pool size, ranging from 3 to 6% of the annual tree stand and understorey uptake. This indicates a more rapid turnover and transport of N, P, and K out of WAN, and suggests that wood ants increase the cycling rate of these nutrients in boreal forests.     

Nest defense behavior against ant attacks in post-emergent colonies of wasp Mischocyttarus cerberus (Hymenoptera, Vespidae)

The primitive eusocial wasp Mischocyttarus cerberus forms colonies of independent foundation, without morphological differentiation among castes. Ants are natural enemies of the social wasps and defending the wasps’ nests involves chemical and active defense strategies. The aims of this work were to verify the kind of defense the wasps use most frequently in post-emergent colonies of M. cerberus. We also observed whether the nest was abandoned during the ant attack and whether any relationship existed between the forms that colony defense took and the number of adults, the number of cells, and the number of immature wasps. The study was carried out on the campus of Universidade Estadual Paulista of Rio Claro, São Paulo, Brazil. The 23 nests under study were mapped weekly, and 68 bioassays were performed by simulating ant attacks against the nests. The results showed that wasps used both active and chemical strategies for nest defense, and the PCA analysis showed that the aggressive behaviors of biting the ant, wings vibrating, gaster hitting, and abdomen pumping were the dominant terms; the PCA correlation values were 2.70, 2.54, 1.64, and 1.63, respectively. The colonies in pre- and post-male substages with few immature wasps and the nests in post-male substage with one adult were more correlated with the nonaggressive behaviors of hiding, staying immobile, and flying; their PCA correlation values were 3.12, 2.56, and 1.77, respectively. These results show that the number of immature wasps is an important factor in the kind and in the intensity of the defense behavior against ant attacks.     

Biological notes on a fungus-growing ant, Trachymyrmex cf. zeteki (Hymenoptera, Formicidae, Attini) attacked by a diverse community of parasitoid wasps (Hymenoptera, Diapriidae)

A number of wasps in the family Diapriidae, subfamily Diapriinae (Proctotrupoidea), are parasitoids that specialize on ant larvae. These wasps are abundant and diverse in the Neotropics, but little is known about their biology. We studied parasitism rates by an array of diapriine wasps that attack the larvae of fungus-growing ants, Trachymyrmex cf. zeteki, in a single population (near Gamboa, Panamá). Relatively little is known about the biology and natural history of these ants, so we also present data on colony size and nest architecture. We excavated 136 colonies in central Panamá from June to September 2006, and 20 nests from July 2009. We reared six wasp morphotypes; two of them in the genus Mimopriella Masner and Garcia, one Oxypria Kieffer, two Szelenyiopria Fabritius and one Acanthopria Ashmead. The mean intensity of larval parasitism per ant colony was 33.9% (2006), and its prevalence across all ant populations was 27.2% (2006 and 2009). Parasitism rates were not positively correlated with host colony size. A single case of super-parasitism was documented in which two Oxypria males were reared from the same host larva.     

How leaf-cutting ants impact forests: drastic nest effects on light environment and plant assemblages

Leaf-cutting ants (Atta spp.) have become a topical issue in Neotropical ecology, particularly because they are reaching hyper-abundance due to escalating levels of fragmentation in recent years. Yet, despite intensive research on their role as dominant herbivores, there is still insufficient documentation on the impacts of their large, long-lived nests on plant assemblage structure and ecosystem functioning. Our study aimed at investigating the magnitude, nature, and spatial extent of nest influence by assessing 11 attributes of ant nest, canopy structure, light environment and sapling assemblage for 20 colonies in four plots along nest-understorey gradients in a large remnant of Atlantic forest. We also monitored the performance of seeds and seedlings of Chrysophyllum viride, an abundant shade-tolerant species. Previously unrecognized canopy gaps above ant nests (0.04–87.9 m²) occurred in 95% of all colonies surveyed. Overall, canopy openness and light availability at least doubled in ant nest plots compared with distant understorey plots. These drastic changes in the light environment paralleled those in plant assemblage: sapling density almost tripled (mean ± SE: 0.42 ± 0.1 saplings m^?2) and sapling species richness doubled (0.16 ± 0.02 species m^?2) in distant plots, as did shade-tolerant species. After a 1-year period, only 33 ± 15.6% of the seeds germinated and all seedlings died on nests, whereas seed germination reached 68 ± 5.1% in distant plots and 66.4 ± 7.6% of their seedlings survived after 12 months. Therefore, plot location was the most significant explanatory variable for predictable and conspicuous changes in the light environment and structure of sapling assemblages. Our findings greatly extend knowledge on the role played by leaf-cutting ants as ecosystem engineers by demonstrating that ant nest-mediated disturbance promotes environmental modifications in tens of meters around nests and is thus, strong enough to drive plant recruitment and consequently alter both the floristic and functional signature of plant assemblages.     

Effect of host species,host nest density and nest size on the occurrence of the shining guest ant <Emphasis Type="Italic">Formicoxenus nitidulus</Emphasis> (Hymenoptera: Formicidae)

Understanding habitat requirements of species is important in conservation. As an obligate ant nest associate, the survival of the globally vulnerable shining guest ant, Formicoxenus nitidulus, is strictly tied to that of its hosts (mound building Formica ants). We investigated how host species, nest density, inter-nest distance and nest mound size relate to the occurrence of F. nitidulus. In total, 166 red wood ant nests were surveyed in SW Finland (120 Formica polyctena, 25 F. rufa, 14 F. aquilonia, 5 F. pratensis, and 2 F. lugubris). Overall, F. nitidulus was found in 60% of the nests. For the actual analysis, only F. polyctena and F. rufa nests were included due to the small number of other nests. F. nitidulus was more likely to be found among F. polyctena than F. rufa. Also, while inter-nest distance was not important, a high nest density, commonly found in polydomous (multi-nest) wood ant colonies, was beneficial for F. nitidulus. The guest ant was also more likely to be found in large host nests than small nests. Thus, our results show that the best habitat for the guest ant is a dense population of host nest mounds with a high proportion of large mounds. Conservation efforts should be directed at keeping the quality of the red wood ant habitats high to preserve their current populations and to increase colonization. This will not only benefit the guest ant, but also a plethora of other species, and help in maintaining the biodiversity of forests.     

Interactions of the ant Crematogaster scutellaris with the fig/fig wasp mutualism

Abstract. 1. A classical example of specialised pollination mutualisms is the relationship between fig trees and their pollinating wasps, in which each partner depends completely on the other for its reproduction; however the fig/fig wasp association is also the target of a great diversity of other species, ranging from specialised parasites to opportunistic foragers, among them ants.
2. The ant community and the sources of ant attraction observed on the Mediterranean fig tree Ficus carica were characterised.
3. A guild of ants attracted by homopterans tended on the plant was distinguished from a second guild composed of two co-dominant ant species ( Crematogaster scutellaris and Pheidole pallidula ) that prey mostly on pollinating wasps, abundant during certain parts of the fig cycle.
4. Foraging workers of C. scutellaris search for prey on the fig inflorescence (syconium), capturing pollinating wasps mostly at the peak of wasp emergence and at a rate estimated to reach 600 prey per day for an entire tree.
5. Detailed study of the predatory sequences displayed under experimental conditions showed that ant workers captured 100% of the pollinating wasps offered, while they captured only 5.5% of the parasitoid wasp specific to the pollinator. The respective impacts of the interaction on ants and on the figs are discussed, as well as several behavioural traits of predation by the dominant ant on pollinators.     

Ant Colonies Prefer Infected over Uninfected Nest Sites

During colony relocation, the selection of a new nest involves exploration and assessment of potential sites followed by colony movement on the basis of a collective decision making process. Hygiene and pathogen load of the potential nest sites are factors worker scouts might evaluate, given the high risk of epidemics in group-living animals. Choosing nest sites free of pathogens is hypothesized to be highly efficient in invasive ants as each of their introduced populations is often an open network of nests exchanging individuals (unicolonial) with frequent relocation into new nest sites and low genetic diversity, likely making these species particularly vulnerable to parasites and diseases. We investigated the nest site preference of the invasive pharaoh ant, Monomorium pharaonis, through binary choice tests between three nest types: nests containing dead nestmates overgrown with sporulating mycelium of the entomopathogenic fungus Metarhizium brunneum (infected nests), nests containing nestmates killed by freezing (uninfected nests), and empty nests. In contrast to the expectation pharaoh ant colonies preferentially (84%) moved into the infected nest when presented with the choice of an infected and an uninfected nest. The ants had an intermediate preference for empty nests. Pharaoh ants display an overall preference for infected nests during colony relocation. While we cannot rule out that the ants are actually manipulated by the pathogen, we propose that this preference might be an adaptive strategy by the host to “immunize” the colony against future exposure to the same pathogenic fungus.     

Arboreal Ant Abundance and Leaf Miner Damage in Coffee Agroecosystems in Mexico

Shaded coffee agroecosystems traditionally have few pest problems potentially due to higher abundance and diversity of predators of herbivores. However, with coffee intensification (e.g., shade tree removal or pruning), some pest problems increase. For example, coffee leaf miner outbreaks have been linked to more intensive management and increased use of agrochemicals. Parasitic wasps control the coffee leaf miner, but few studies have examined the role of predators, such as ants, that are abundant and diverse in coffee plantations. Here, we examine linkages between arboreal ant communities and coffee leaf miner incidence in a coffee plantation in Mexico. We examined relationships between incidence and severity of leaf miner attack and: (1) variation in canopy cover, tree density, tree diversity, and relative abundance of Inga spp. shade trees; (2) presence of Azteca instabilis, an arboreal canopy dominant ant; and (3) the number of arboreal twig‐nesting ant species and nests in coffee plants. Differences in vegetation characteristics in study plots did not correlate with leaf miner damage perhaps because environmental factors act on pest populations at a larger spatial scale. Further, presence of A. instabilis did not influence presence or severity of leaf miner damage. The proportion of leaves with leaf miner damage was significantly lower where abundance of twig‐nesting ants was higher but not where twig‐nesting ant richness was higher. These results indicate that abundance of twig‐nesting ants in shaded coffee plantations may contribute to maintenance of low leaf miner populations and that ants provide important ecosystem services in coffee agroecosystems.     

Ant behaviour and seed morphology: a missing link of myrmecochory

Seed dispersal by ants (myrmecochory) is mediated by the presence of a lipid-rich appendage (elaiosome) on the seed that induces a variety of ants to collect the diaspores. When seeds mature or fall onto the ground, these ant species transport them to their nest. After eating the elaiosome, the seed is discarded in nest galleries or outside, in the midden or farther away, where seeds can potentially germinate. The final location of seeds with their elaiosomes removed was evaluated to assess the importance of possible handles (structures that ants can grasp to carry) in transporting ants during re-dispersal experiments of seeds from nests of six species of ants. The results indicate that seeds remained within the nest because the ants were not able to transport them out of the nest. As a consequence of the elaiosome being removed, small ant species could not take Euphorbia characias seeds out of their nests. Only large ant species could remove E. characias seeds from their nests. Attaching an artificial handle to E. characias seeds allowed small ant species to redistribute the seeds from their nests. On the other hand, Rhamnus alaternus seeds that have a natural handle after the elaiosome removal were removed from the nests by both groups of ant species. If a seed has an element that acts as a handle, it will eventually get taken out of the nest. The ants’ size and their mandible gap can determine the outcome of the interaction (i.e. the pattern of the final seed shadow) and as a consequence, could influence the events that take place after the dispersal process.     

Comparative characteristics of nests of Vespula germanica (F.) and Vespula vulgaris (L.) (Hymenoptera: Vespinae) from Christchurch City,New Zealand

Abstract

From July 1987 to June 1989, 141 nests and 365 nests, respectively, of the adventive social vespines the German wasp, Vespula germanica (F.), and the common wasp V. vulgaris (L.), from the City of Christchurch, New Zealand, were evaluated for seven major characteristics.

There were few, if any, differences in nest sites, and from spring to early summer in nest traffic, nest size, and numbers of combs. However, Getman wasps showed no preference for direction of nest entrances, while common wasp nests were more numerous in areas most exposed to the morning sun. Some Getman wasp nests survived the winter and began producing new worker cells by late June, but all common wasp nests died by June. Because common wasp nests have been reported surviving the winter in beech forest, which produces honey dew, and honey dew is not available in Christchurch City, common wasp nest survival over winter may be more dependent upon carbohydrates and/or prey dependent upon carbohydrates than the survival of German wasp nests. The presence of large, expanding wasp nests from early spring must impose localised predation pressures virtually unknown in the Northern Hemisphere where overwintering nests are rare. Wasp population dynamics, and impacts of wasps on fauna, are likely to vary between different geographical areas of New Zealand, but as targets for biological control, the two wasp species can generally be considered to be quite similar.     

Disentangling the Diversity of Arboreal Ant Communities in Tropical Forest Trees

Tropical canopies are known for their high abundance and diversity of ants. However, the factors which enable coexistence of so many species in trees, and in particular, the role of foragers in determining local diversity, are not well understood. We censused nesting and foraging arboreal ant communities in two 0.32 ha plots of primary and secondary lowland rainforest in New Guinea and explored their species diversity and composition. Null models were used to test if the records of species foraging (but not nesting) in a tree were dependent on the spatial distribution of nests in surrounding trees. In total, 102 ant species from 389 trees occurred in the primary plot compared with only 50 species from 295 trees in the secondary forest plot. However, there was only a small difference in mean ant richness per tree between primary and secondary forest (3.8 and 3.3 sp. respectively) and considerably lower richness per tree was found only when nests were considered (1.5 sp. in both forests). About half of foraging individuals collected in a tree belonged to species which were not nesting in that tree. Null models showed that the ants foraging but not nesting in a tree are more likely to nest in nearby trees than would be expected at random. The effects of both forest stage and tree size traits were similar regardless of whether only foragers, only nests, or both datasets combined were considered. However, relative abundance distributions of species differed between foraging and nesting communities. The primary forest plot was dominated by native ant species, whereas invasive species were common in secondary forest. This study demonstrates the high contribution of foragers to arboreal ant diversity, indicating an important role of connectivity between trees, and also highlights the importance of primary vegetation for the conservation of native ant communities.     

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.     

Utilization of harvester ant nest sites by Persian goitered gazelle in steppes of central Iran

Ants are among the most important elements in many ecosystems and known as famous ecosystem engineers. By changing physical and chemical properties of soil, ants may provide suitable habitats for other species. Based on previous observations, we hypothesized that Persian goitered gazelles (Gazella subgutturosa subgutturosa) exhibit a preference for utilizing sites close to seed harvester ant (Messor spp.) nests. We tested our hypothesis by (1) mapping the occurrence of harvester ant nests and aggregated gazelle pellet groups along 31 strip transects, (2) monitoring pellet group accumulation bimonthly at 56 pairs of permanent plots established on ant nests and at adjacent control sites for a complete year, and (3) comparing vegetation and soil parameters between ant nest sites used by gazelles and paired control plots without ant nests. Although the area of Messor spp. nest sites covered only about 0.29% of the sampled transects, 84% of the gazelle pellet group aggregation sites were positioned upon ant nests, suggesting that gazelles actively selected Messor spp. nest sites. Pair-wise comparisons between ant nest plots and paired control plots also confirmed higher use of ant nest sites by gazelles compared to sites without ant nests in all time periods. Percent soil organic matter, percent cover of gravel, and annual herb vegetation significantly differed between ant nest and paired control plots in all the vegetation communities. We suggest that the alterations brought about by harvester ants on soil and vegetation make these sites attractive to gazelles. Gazelle territoriality behaviour and use of ant nests as bedding sites may be the reasons for selection of ant nest sites by gazelles.