One of the most successful invasive species is the common wasp, Vespula vulgaris. We recently reported how foragers of this species have adopted previously unknown interference behavior when competing for food with native ants. Picking their opponents up in their mandibles, flying backward and dropping them some distance away from the disputed resource, wasps were shown to efficiently deal with a yet aggressive competitor and to modulate this behavior according to circumstances. Here we further discuss the nature and functioning of this unusual strategy. We first highlight the questions this interaction raises regarding the competitive advantages offered by asymmetries in body size and flight ability. Then, we argue that this study system illustrates the important role of behavioral plasticity in biological invasions; not only in the success of invaders but also in the ability of native species to coexist with these invaders.
biological invasions; interference behavior; Prolasius advenus; social insects; Vespula vulgaris
Throughout lowland Amazonia, arboreal ants collect seeds of specific plants and cultivate them in nutrient-rich nests, forming diverse yet obligate and species-specific symbioses called Neotropical ant-gardens (AGs). The ants depend on their symbiotic plants for nest stability, and the plants depend on AGs for substrate and nutrients. Although the AGs are limited to specific participants, it is unknown at what stage specificity arises, and seed fate pathways in AG epiphytes are undocumented. Here we examine the specificity of the ant-seed interaction by comparing the ant community observed at general food baits to ants attracted to and removing seeds of the AG plant Peperomia macrostachya. We also compare seed removal rates under treatments that excluded vertebrates, arthropods, or both. In the bait study, only three of 70 ant species collected P. macrostachya seeds, and 84% of observed seed removal by ants was attributed to the AG ant Camponotus femoratus. In the exclusion experiment, arthropod exclusion significantly reduced seed removal rates, but vertebrate exclusion did not. We provide the most extensive empirical evidence of species specificity in the AG mutualism and begin to quantify factors that affect seed fate in order to understand conditions that favor its departure from the typical diffuse model of plant-animal mutualism.
Social insect colonies are like fortresses, well protected and rich in shared stored resources. This makes them ideal targets for exploitation by predators, parasites and competitors. Colonies of Myrmica rubra ants are sometimes exploited by the parasitic butterfly Maculinea alcon. Maculinea alcon gains access to the ants' nests by mimicking their cuticular hydrocarbon recognition cues, which allows the parasites to blend in with their host ants. Myrmica rubra may be particularly susceptible to exploitation in this fashion as it has large, polydomous colonies with many queens and a very viscous population structure. We studied the mutual aggressive behaviour of My. rubra colonies based on predictions for recognition effectiveness. Three hypotheses were tested: first, that aggression increases with distance (geographical, genetic and chemical); second, that the more queens present in a colony and therefore the less-related workers within a colony, the less aggressively they will behave; and that colonies facing parasitism will be more aggressive than colonies experiencing less parasite pressure. Our results confirm all these predictions, supporting flexible aggression behaviour in Myrmica ants depending on context.
aggression; defence; social parasite; Maculinea
Argentine ants (Linepithema humile) live in groups of nests connected by trails to each other and to stable food sources. In a field study, we investigated whether some ants recruit directly from established, persistent trails to food sources, thus accelerating food collection. Our results indicate that Argentine ants recruit nestmates to food directly from persistent trails, and that the exponential increase in the arrival rate of ants at baits is faster than would be possible if recruited ants traveled from distant nests. Once ants find a new food source, they walk back and forth between the bait and sometimes share food by trophallaxis with nestmates on the trail. Recruiting ants from nearby persistent trails creates a dynamic circuit, like those found in other distributed systems, which facilitates a quick response to changes in available resources.
The Argentine ant, Linepithema humile, is a widespread invasive ant species that has successfully established in nearly all continents across the globe. Argentine ants are characterised by a social structure known as unicoloniality, where territorial boundaries between nests are absent and intraspecific aggression is rare. This is particularly pronounced in introduced populations and results in the formation of large and spatially expansive supercolonies. Although it is amongst the most well studied of invasive ants, very little work has been done on this ant in South Africa. In this first study, we investigate the population structure of Argentine ants in South Africa. We use behavioural (aggression tests) and chemical (CHC) approaches to investigate the population structure of Argentine ants within the Western Cape, identify the number of supercolonies and infer number of introductions.
Both the aggression assays and chemical data revealed that the Western Cape Argentine ant population can be divided into two behaviourally and chemically distinct supercolonies. Intraspecific aggression was evident between the two supercolonies of Argentine ants with ants able to discriminate among conspecific non-nestmates. This discrimination is linked to the divergence in cuticular hydrocarbon profiles of ants originating from the two supercolonies.
The presence of these two distinct supercolonies is suggestive of at least two independent introductions of this ant within the Western Cape. Moreover, the pattern of colonisation observed in this study, with the two colonies interspersed, is in agreement with global patterns of Argentine ant invasions. Our findings are of interest because recent studies show that Argentine ants from South Africa are different from those identified in other introduced ranges and therefore provide an opportunity to further understand factors that determine the distributional and spread patterns of Argentine ant supercolonies.
Vespula germanica (F.) (Hymenoptera: Vespidae) is a highly invasive social wasp that exhibits a rich behavioral repertoire in which learning and memory play a fundamental role in foraging. The learning abilities of these wasps were analyzed while relocating a food source and whether V. germanica foragers are capable of discriminating between different orientation patterns and generalizing their choice to a new pattern. Foraging wasps were trained to associate two different stripe orientation patterns with their respective food locations. Their response to a novel configuration that maintained the orientation of one of the learned patterns but differed in other aspects (e.g. width of stripes) was then evaluated. The results support the hypothesis that V. germanica wasps are able to associate a particular oriented pattern with the location of a feeder and to generalize their choice to a new pattern, which differed in quality, but presented the same orientation.
learning; social wasps; visual patterns
High recombination rates have previously been detected in two groups of eusocial insects; honeybees and ants. In this study we estimate recombination rate in a eusocial wasp Vespula vulgaris that represents a third phylogenetic lineage within eusocial hymenopterans.
A genetic linkage map of V. vulgaris based on 210 markers shows that the total map length is 2129 cM and the recombination rate is 9.7 cM/Mb (or 103 kb/cM). The present estimate in V. vulgaris is somewhat smaller than in the honeybee Apis mellifera and intermediate between the estimates from two ant species (Acromyrmex echinatior, Pogonomyrmex rugosus). Altogether, the estimates from these eusocial species are higher than in any other insect reported so far.
The four species (V. vulgaris, A. mellifera, A. echinatior, P. rugosus) are characterized by advanced eusociality with large colonies, clear queen-worker dimorphism and well developed task specialization. They also have colonies with a single, normally multiply inseminated (polyandrous) queen. Benefits of genotypic diversity within colonies (e.g. through improved task specialization or pathogen and parasite resistance) may have selected for both polyandry and high recombination rate in such advanced eusocial insects.
A comparison of the thermoregulation of water foraging wasps (Vespula vulgaris, Polistes dominulus) under special consideration of ambient temperature and solar radiation was conducted. The body surface temperature of living and dead wasps was measured by infrared thermography under natural conditions in their environment without disturbing the insects’ behaviour. The body temperature of both of them was positively correlated with Ta and solar radiation. At moderate Ta (22–28 °C) the regression lines revealed mean thorax temperatures (Tth) of 35.5–37.5 °C in Vespula, and of 28.6–33.7 °C in Polistes. At high Ta (30–39 °C) Tth was 37.2–40.6 °C in Vespula and 37.0–40.8 °C in Polistes. The thorax temperature excess (Tth–Ta) increased at moderate Ta by 1.9 °C (Vespula) and 4.4 °C (Polistes) per kW−1 m−2. At high Ta it increased by 4.0 °C per kW−1 m−2 in both wasps. A comparison of the living water foraging Vespula and Polistes with dead wasps revealed a great difference in their thermoregulatory behaviour. At moderate Ta (22–28 °C) Vespula exhibited distinct endothermy in contrast to Polistes, which showed only a weak endothermic activity. At high Ta (30–39 °C) Vespula reduced their active heat production, and Polistes were always ectothermic. Both species exhibited an increasing cooling effort with increasing insolation and ambient temperature.
Wasp; Thermoregulation; Foraging; Thermography; Water
Desert ants, Cataglyphis fortis, are equipped with remarkable skills that enable them to navigate efficiently. When travelling between the nest and a previously visited feeding site, they perform path integration (PI), but pinpoint the nest or feeder by following odour plumes. Homing ants respond to nest plumes only when the path integrator indicates that they are near home. This is crucial, as homing ants often pass through plumes emanating from foreign nests and do not discriminate between the plume of their own and that of a foreign nest, but should absolutely avoid entering a wrong nest. Their behaviour towards food odours differs greatly. Here, we show that in ants on the way to food, olfactory information outweighs PI information. Although PI guides ants back to a learned feeder, the ants respond to food odours independently of whether or not they are close to the learned feeding site. This ability is beneficial, as new food sources—unlike foreign nests—never pose a threat but enable ants to shorten distances travelled while foraging. While it has been shown that navigating C. fortis ants rely strongly on PI, we report here that the ants retained the necessary flexibility in the use of PI.
Cataglyphis; path integration; odour plume; homing; foraging; navigation
Ants and termites are the most abundant animals on earth. Their ecological success is attributed to their social life. They live in colonies consisting of few reproducing individuals, while the large majority of colony members (workers/soldiers) forego reproduction at least temporarilly. Despite their apparent resemblance in social organisation, both groups evolved social life independently. Termites are basically social cockroaches, while ants evolved from predatory wasps. In this review, I will concentrate on termites with an ancestral life type, the wood-dwelling termites, to compare them with ants. Their different ancestries provided both groups with different life history pre-adaptations for social evolution. Like their closest relatives, the woodroaches, wood-dwelling termites live inside their food, a piece of wood. Thus, intensive costly food provisioning of their young is not necessary, especially as young instars are rather independent due to their hemimetabolous development. In contrast, ants are progressive food provisioners which have to care intensively for their helpless brood. Corresponding to the precocial – altricial analogy, helping by workers is selected in ants, while new evidence suggests that wood-dwelling termite workers are less engaged in brood care. Rather they seem to stay in the nest because there is generally low selection for dispersal. The nest presents a safe haven with no local resource competition as long as food is abundant (which is generally the case), while founding a new colony is very risky. Despite these differences between ants and termites, their common dwelling life style resulted in convergent evolution, especially winglessness, that probably accounts for the striking similarity between both groups. In ants, all workers are wingless and winglessness in sexuals evolved in several taxa as a derived trait. In wood-dwelling termites, workers are by default wingless as they are immatures. These immatures can develop into winged sexuals that disperse and found a new nest or into neotenic replacement reproductives that inherit the natal colony. Depending on the worker instar from which the latter develop, the neotenic reproductives are either apterous or brachypterous, but never winged. I propose that this wing polyphenism might present a basis for the evolution of social life in termites.
Plant-ants live in a mutualistic association with host plants known as “myrmecophytes” that provide them with a nesting place and sometimes with extra-floral nectar (EFN) and/or food bodies (FBs); the ants can also attend sap-sucking Hemiptera for their honeydew. In return, plant-ants, like most other arboreal ants, protect their host plants from defoliators. To satisfy their nitrogen requirements, however, some have optimized their ability to capture prey in the restricted environment represented by the crowns of trees by using elaborate hunting techniques. In this study, we investigated the predatory behavior of the ant Azteca andreae which is associated with the myrmecophyte Cecropia obtusa. We noted that up to 8350 ant workers per tree hide side-by-side beneath the leaf margins of their host plant with their mandibles open, waiting for insects to alight. The latter are immediately seized by their extremities, and then spread-eagled; nestmates are recruited to help stretch, carve up and transport prey. This group ambush hunting technique is particularly effective when the underside of the leaves is downy, as is the case for C. obtusa. In this case, the hook-shaped claws of the A. andreae workers and the velvet-like structure of the underside of the leaves combine to act like natural Velcro® that is reinforced by the group ambush strategy of the workers, allowing them to capture prey of up to 13,350 times the mean weight of a single worker.
In ant–plant symbioses, plants provide symbiotic ants with food and specialized nesting cavities (called domatia). In many ant–plant symbioses, a fungal patch grows within each domatium. The symbiotic nature of the fungal association has been shown in the ant-plant Leonardoxa africana and its protective mutualist ant Petalomyrmex phylax. To decipher trophic fluxes among the three partners, food enriched in 13C and 15N was given to the ants and tracked in the different parts of the symbiosis up to 660 days later. The plant received a small, but significant, amount of nitrogen from the ants. However, the ants fed more intensively the fungus. The pattern of isotope enrichment in the system indicated an ant behaviour that functions specifically to feed the fungus. After 660 days, the introduced nitrogen was still present in the system and homogeneously distributed among ant, plant and fungal compartments, indicating efficient recycling within the symbiosis. Another experiment showed that the plant surface absorbed nutrients (in the form of simple molecules) whether or not it is coated by fungus. Our study provides arguments for a mutualistic status of the fungal associate and a framework for investigating the previously unsuspected complexity of food webs in ant–plant mutualisms.
mutualism; symbiosis; Myrmecophyte; nutrient transfer; stable isotopes; Leonardoxa
The ancient mutualism between fungus-growing ants and the fungi they cultivate for food is a textbook example of symbiosis. Fungus-growing ants' ability to cultivate fungi depends on protection of the garden from the aggressive microbes associated with the substrate added to the garden as well as from the specialized virulent garden parasite Escovopsis. We examined ants' ability to remove alien microbes physically by infecting Atta colombica gardens with the generalist pathogen Trichoderma viride and the specialist pathogen Escovopsis. The ants sanitized the garden using two main behaviours: grooming of alien spores from the garden (fungus grooming) and removal of infected garden substrate (weeding). Unlike previously described hygienic behaviours (e.g. licking and self-grooming), fungus-grooming and garden-removal behaviours are specific responses to the presence of fungal pathogens. In the presence of pathogens, they are the primary activities performed by workers, but they are uncommon in uninfected gardens. In fact, workers rapidly eliminate Trichoderma from their gardens by fungus grooming and weeding, suggesting that these behaviours are the primary method of garden defence against generalist pathogens. The same sanitary behaviours were performed in response to the presence of the specialist pathogen Escovopsis. However, the intensity and duration of these behaviours were much greater in this treatment. Despite the increased effort, the ants were unable to eliminate Escovopsis from their gardens, suggesting that this specialized pathogen has evolved counter-adaptations in order to overcome the sanitary defences of the ants.
Ants form highly social and cooperative colonies that compete, and often fight, against other such colonies, both intra- and interspecifically. Some invasive ants take sociality to an extreme, forming geographically massive 'supercolonies' across thousands of kilometres. The success of social insects generally, as well as invasive ants in particular, stems from the sophisticated mechanisms used to accurately and precisely distinguish colonymates from non-colonymates. Surprisingly, however, the specific chemicals used for this recognition are virtually undescribed.
Here, we report the discovery, chemical synthesis and behavioural testing of the colonymate recognition cues used by the widespread and invasive Argentine ant (Linepithema humile). By synthesizing pure versions of these chemicals in the laboratory and testing them in behavioural assays, we show that these compounds trigger aggression among normally amicable nestmates, but control hydrocarbons do not. Furthermore, behavioural testing across multiple different supercolonies reveals that the reaction to individual compounds varies from colony to colony -- the expected reaction to true colony recognition labels. Our results also show that both quantitative and qualitative changes to cuticular hydrocarbon profiles can trigger aggression among nestmates. These data point the way for the development of new environmentally-friendly control strategies based on the species-specific manipulation of aggressive behaviour.
Overall, our findings reveal the identity of specific chemicals used for colonymate recognition by the invasive Argentine ants. Although the particular chemicals used by other ants may differ, the patterns reported here are likely to be true for ants generally. As almost all invasive ants display widespread unicoloniality in their introduced ranges, our findings are particularly relevant for our understanding of the biology of these damaging invaders.
The Argentine ant (Linepithema humile) is recognized as one of the world's most damaging invasive species. One reason for the ecological dominance of introduced Argentine ant populations is their ability to dominate food and habitat resources through the rapid mobilization and recruitment of thousands of workers. More than 30 years ago, studies showed that (Z)-9-hexadecenal strongly attracted Argentine ant workers in a multi-choice olfactometer, suggesting that (Z)-9-hexadecenal might be the trail pheromone, or a component of a trail pheromone mixture. Since then, numerous studies have considered (Z)-9-hexadecenal as the key component of the Argentine ant trails. Here, we report the first chemical analyses of the trails laid by living Argentine ants and find that (Z)-9-hexadecenal is not present in a detectible quantity. Instead, two iridoids, dolichodial and iridomyrmecin, appear to be the primary chemical constituents of the trails. Laboratory choice tests confirmed that Argentine ants were attracted to artificial trails comprised of these two chemicals significantly more often than control trails. Although (Z)-9-hexadecenal was not detected in natural trails, supplementation of artificial dolichodial+iridomyrmecin trails with an extremely low concentraion of (Z)-9-hexadecenal did increase the efficacy of the trail-following behavior. In stark contrast with previous dogma, our study suggests that dolichodial and iridomyrmecin are major components of the Argentine ant trail pheromone. (Z)-9-hexadecenal may act in an additive manner with these iridoids, but it does not occur in detectable quantities in Argentine ant recruitment trails.
Although mutualism between ants and honeydew-producing hemipterans has been extensively recognized in ecosystem biology, however few attempts to test the hypothesis that mutualism between two alien species leads to the facilitation of the invasion process. To address this problem, we focus on the conditional mutualism between S. invicta and P. solenopsis by field investigations and indoor experiments. In the laboratory, ant colony growth increased significantly when ants had access to P. solenopsis and animal-based food. Honeydew produced by P. solenopsis also improved the survival of ant workers. In the field, colony density of P. solenopsis was significantly greater on plots with ants than on plots without ants. The number of mealybug mummies on plants without fire ants was almost three times that of plants with fire ants, indicating a strong effect of fire ants on mealybug survival. In addition, the presence of S. invicta successfully contributed to the spread of P. solenopsis. The quantity of honeydew consumption by S. invicta was significantly greater than that of a presumptive native ant, Tapinoma melanocephalum. When compared with the case without ant tending, mealybugs tended by ants matured earlier and their lifespan and reproduction increased. T. melanocephalum workers arrived at honeydew more quickly than S. invicta workers, while the number of foraging S. invicta workers on plants steadily increased, eventually exceeding that number of T. melanocephalum foragers. Overall, these results suggest that the conditional mutualism between S. invicta and P. solenopsis facilitates population growth and fitness of both species. S. invicta tends to acquire much more honeydew and drive away native ants, promoting their predominance. These results suggest that the higher foraging tempo of S. invicta may provide more effective protection of P. solenopsis than native ants. Thus mutualism between these two alien species may facilitate the invasion success of both species.
Hornets, the largest social wasps, have a reputation of being facultatively nocturnal. Here we confirm flight activity of hornet workers in dim twilight. We studied the eyes and ocelli of European hornets (Vespa crabro) and common wasps (Vespula vulgaris) with the goal to find the optical and anatomical adaptations that enable them to fly in dim light. Adaptations described for obligately nocturnal hymenoptera such as the bees Xylocopa tranquebarica and Megalopta genalis and the wasp Apoica pallens include large ocelli and compound eyes with wide rhabdoms and large facet lenses. Interestingly, we did not find any such adaptations in hornet eyes or ocelli. On the contrary, their eyes are even less sensitive than those of the obligately diurnal common wasps. Therefore we conclude that hornets, like several facultatively nocturnal bee species such as Apis mellifera adansonii, A. dorsata and X. tenuiscapa are capable of seeing in dim light simply due to the large body and thus eye size. We propose that neural pooling strategies and behavioural adaptations precede anatomical adaptations in the eyes and ocelli when insects with apposition compound eyes turn to dim light activity.
Ant-plants provide food and nesting space (domatia) for ants that protect them against herbivores. These mutualisms are often very specific and are usually considered as bipartite, or tripartite when ants use hemipterans as trophobionts. However, fungi growing inside domatia have been recorded by a few authors. Here we report on their occurrence on additional ant-plants from Africa, Asia and South America. We demonstrated the symbiotic nature of the relationship between the plant, the ant and the fungus in the model plant Leonardoxa africana africana and its mutualistic ant Petalomyrmex phylax. Moreover, data suggest the ant-fungus relationship is mutualistic. Here we discuss the most probable role of the fungus and the potential implications on the understanding of nutritional ecology of ant-plant symbioses. The fungus is also associated with the presence of nematodes and bacteria. Many ant-plant symbioses previously considered to be bipartite will soon likely prove to be multipartite symbiotic communities.
symbiosis; mutualism; ant-plant; fungi; community ecology; Leonardoxa; coevolution; nematodes
Termites are instantly recognizable mound-builders and house-eaters: their complex social lifestyles have made them incredibly successful throughout the tropics. Although known as ‘white ants’, they are not ants and their relationships with other insects remain unclear. Our molecular phylogenetic analyses, the most comprehensive yet attempted, show that termites are social cockroaches, no longer meriting being classified as a separate order (Isoptera) from the cockroaches (Blattodea). Instead, we propose that they should be treated as a family (Termitidae) of cockroaches. It is surprising to find that a group of wood-feeding cockroaches has evolved full sociality, as other ecologically dominant fully social insects (e.g. ants, social bees and social wasps) have evolved from solitary predatory wasps.
insect systematics; phylogenetics; social evolution; classification
Obligate mating of females (queens) with multiple males has evolved only rarely in social Hymenoptera (ants, social bees, social wasps) and for reasons that are fundamentally different from those underlying multiple mating in other animals. The monophyletic tribe of ('attine') fungus-growing ants is known to include evolutionarily derived genera with obligate multiple mating (the Acromyrmex and Atta leafcutter ants) as well as phylogenetically basal genera with exclusively single mating (e.g. Apterostigma, Cyphomyrmex, Myrmicocrypta). All attine genera share the unique characteristic of obligate dependence on symbiotic fungus gardens for food, but the sophistication of this symbiosis differs considerably across genera. The lower attine genera generally have small, short-lived colonies and relatively non-specialized fungal symbionts (capable of living independently of their ant hosts), whereas the four evolutionarily derived higher attine genera have highly specialized, long-term clonal symbionts. In this paper, we investigate whether the transition from single to multiple mating occurred relatively recently in the evolution of the attine ants, in conjunction with the novel herbivorous 'leafcutter' niche acquired by the common ancestor of Acromyrmex and Atta, or earlier, at the transition to rearing specialized long-term clonal fungi in the common ancestor of the larger group of higher attines that also includes the genera Trachymyrmex and Sericomyrmex. We use DNA microsatellite analysis to provide unambiguous evidence for a single, late and abrupt evolutionary transition from exclusively single to obligatory multiple mating. This transition is historically correlated with other evolutionary innovations, including the extensive use of fresh vegetation as substrate for the fungus garden, a massive increase in mature colony size and morphological differentiation of the worker caste.
Interactions between ants and honeydew-producing hemipteran insects are abundant and widespread in arthropod food webs, yet their ecological consequences are very poorly known. Ant–hemipteran interactions have potentially broad ecological effects, because the presence of honeydew-producing hemipterans dramatically alters the abundance and predatory behaviour of ants on plants. We review several studies that investigate the consequences of ant–hemipteran interactions as ‘keystone interactions’ on arthropod communities and their host plants. Ant–hemipteran interactions have mostly negative effects on the local abundance and species richness of several guilds of herbivores and predators. In contrast, out of the 30 studies that document the effects of ant–hemipteran interactions on plants, the majority (73%) shows that plants actually benefit indirectly from these interactions. In these studies, increased predation or harassment of other, more damaging, herbivores by hemipteran-tending ants resulted in decreased plant damage and/or increased plant growth and reproduction. The ecological consequences of mutualistic interactions between honeydew-producing hemipterans and invasive ants relative to native ants have rarely been studied, but they may be of particular importance owing to the greater abundance, aggressiveness and extreme omnivory of invasive ants. We argue that ant–hemipteran interactions are largely overlooked and underappreciated interspecific interactions that have strong and pervasive effects on the communities in which they are embedded.
ants; hemipterans; mutualism; indirect effects; tritrophic interactions; conditionality
Environmental conditions and physical constraints both influence an animal's behavior. We investigate whether behavioral variation among colonies of the black harvester ant, Messor andrei, remains consistent across foraging and disturbance situations and ask whether consistent colony behavior is affected by nest site and weather. We examined variation among colonies in responsiveness to food baits and to disturbance, measured as a change in numbers of active ants, and in the speed with which colonies retrieved food and removed debris. Colonies differed consistently, across foraging and disturbance situations, in both responsiveness and speed. Increased activity in response to food was associated with a smaller decrease in response to alarm. Speed of retrieving food was correlated with speed of removing debris. In all colonies, speed was greater in dry conditions, reducing the amount of time ants spent outside the nest. While a colony occupied a certain nest site, its responsiveness was consistent in both foraging and disturbance situations, suggesting that nest structure influences colony personality.
behavioral syndromes; collective behavior; harvester ant; Messor andrei; nest structure; personality; plasticity; social insects; temperament
Understanding how species will respond to global change depends on our ability to distinguish generalities from idiosyncrasies. For diverse, but poorly known taxa, such as insects, species traits may provide a short-cut to predicting species turnover. We tested whether ant traits respond consistently to habitat complexity across geographically independent ant assemblages, using an experimental approach and baits. We repeated our study in six paired simple and complex habitats on three continents with distinct ant faunas. We also compared traits amongst ants with different foraging strategies. We hypothesised that ants would be larger, broader, have longer legs and more dorsally positioned eyes in simpler habitats. In agreement with predictions, ants had longer femurs and dorsally positioned eyes in simple habitats. This pattern was most pronounced for ants that discovered resources. Body size and pronotum width responded as predicted for experimental treatments, but were inconsistent across continents. Monopolising ants were smaller, with shorter femurs than those that occupied or discovered resources. Consistent responses for several traits suggest that many, but not all, aspects of morphology respond predictably to habitat complexity, and that foraging strategy is linked with morphology. Some traits thus have the potential to be used to predict the direction of species turnover, changes in foraging strategy and, potentially, evolution in response to changes in habitat structure.
Foraging workers of grass-cutting ants (Atta vollenweideri) regularly carry grass fragments larger than their own body. Fragment length has been shown to influence the ants’ running speed and thereby the colony’s food intake rate. We investigated whether and how grass-cutting ants maintain stability when carrying fragments of two different lengths but identical mass.
Ants carried all fragments in an upright, backwards-tilted position, but held long fragments more vertically than short ones. All carrying ants used an alternating tripod gait, where mechanical stability was increased by overlapping stance phases of consecutive steps. The overlap was greatest for ants carrying long fragments, resulting in more legs contacting the ground simultaneously. For all ants, the projection of the total centre of mass (ant and fragment) was often outside the supporting tripod, i.e. the three feet that would be in stance for a non-overlapping tripod gait. Stability was only achieved through additional legs in ground contact. Tripod stability (quantified as the minimum distance of the centre of mass to the edge of the supporting tripod) was significantly smaller for ants with long fragments. Here, tripod stability was lowest at the beginning of each step, when the center of mass was near the posterior margin of the supporting tripod. By contrast, tripod stability was lowest at the end of each step for ants carrying short fragments. Consistently, ants with long fragments mainly fell backwards, whereas ants carrying short fragments mainly fell forwards or to the side. Assuming that transporting ants adjust neither the fragment angle nor the gait, they would be less stable and more likely to fall over.
In grass-cutting ants, the need to maintain static stability when carrying long grass fragments has led to multiple kinematic adjustments at the expense of a reduced material transport rate.
Homing by the nocturnal Namib Desert spider Leucorchestris arenicola (Araneae: Sparassidae) is comparable to homing in diurnal bees, wasps and ants in terms of path length and layout. The spiders' homing is based on vision but their basic navigational strategy is unclear. Diurnal homing insects use memorised views of their home in snapshot matching strategies. The insects learn the visual scenery identifying their nest location during learning flights (e.g. bees and wasps) or walks (ants). These learning flights and walks are stereotyped movement patterns clearly different from other movement behaviours. If the visual homing of L. arenicola is also based on an image matching strategy they are likely to exhibit learning walks similar to diurnal insects. To explore this possibility we recorded departures of spiders from a new burrow in an unfamiliar area with infrared cameras and analysed their paths using computer tracking techniques. We found that L. arenicola performs distinct stereotyped movement patterns during the first part of their departures in an unfamiliar area and that they seem to learn the appearance of their home during these movement patterns. We conclude that the spiders perform learning walks and this strongly suggests that L. arenicola uses a visual memory of the burrow location when homing.