Nestmate recognition in Camponotus floridanus callow worker ants: are sisters or nestmates recognized?

Callow workers of the Florida carpenter ant, Camponotus floridanus Buckley, reared with their sisters displayed more aggressive acts towards unrelated unfamiliar foragers than towards related unfamiliar foragers. When callows were reared with foreign nurses they were more aggressive towards their own unfamiliar sisters than towards the unfamiliar sisters of their nurses. Thus callow workers recognized the sisters of the workers they were reared with as nestmates. This ability was not dependent on whether or not callows received cues from older ants during their emergence. Similarly, a forager was less aggressive towards unfamiliar callows reared with its sisters than towards those reared with nurses unrelated to itself. In addition, foragers were less aggressive towards their callow sisters than towards unrelated callows, regardless of the relatedness between the foragers and the rearing nurses.     

Seed handling behaviours of native and invasive seed‐dispersing ants differentially influence seedling emergence in an introduced plant

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Similar yet different: differential response of a praying mantis to ant‐mimicking spiders

Batesian mimics typically dupe visual predators by resembling noxious or deadly model species. Ants are unpalatable and dangerous to many arthropod taxa, and are popular invertebrate models in mimicry studies. Ant mimicry by spiders, especially jumping spiders, has been studied and researchers have examined whether visual predators can distinguish between the ant model, spider mimic and spider non‐mimics. Tropical habitats harbour a diverse community of ants, their mimics and predators. In one such tripartite mimicry system, we investigated the response of an invertebrate visual predator, the ant‐mimicking praying mantis (Euantissa pulchra), to two related ant‐mimicking spider prey of the genus Myrmarachne, each closely mimicking its model ant species. We found that weaver ants (Oecophylla smaragdina) were much more aggressive than carpenter ants (Camponotus sericeus) towards the mantis. Additionally, mantids exhibited the same aversive response towards ants and their mimics. More importantly, mantids approached carpenter ant‐mimicking spiders significantly more than often that they approached weaver ant‐mimicking spiders. Thus, in this study, we show that an invertebrate predator, the praying mantis, can indeed discriminate between two closely related mimetic prey. The exact mechanism of the discrimination remains to be tested, but it is likely to depend on the level of mimetic accuracy by the spiders and on the aggressiveness of the ant model organism.     

Temporal polyethism, life expectancy, and entropy of workers of the ant Ectatomma vizottoi Almeida, 1987 (Formicidae: Ectatomminae)

We investigated the changes in the behavioral repertoire over the course of life and determined the life expectancy and entropy of workers of the ant Ectatomma vizottoi. Newly emerged ants were individually marked with model airplane paint for observation of behaviors and determination of the age and life expectancy. Ants were divided into two groups: young and old workers. The 36 behaviors observed were divided into eight categories. Workers exhibit a clear division of tasks throughout their lives, with young workers performing more tasks inside the colony and old workers, outside, unlike species that have small colonies. This species also exhibits an intermediate life expectancy compared to workers of other species that are also intermediary in size. This supports the hypothesis of a relationship between size and maximum life expectancy, but it also suggests that other factors may also be acting in concert. Entropy value shows a high mortality rate during the first life intervals.     

Early Learning and the Recognition of Conspecific Cocoons by Carpenter Ants (Camponotus spp.)

Groups of newly-eclosed workers of two carpenter ant species (Camponotus floridanus and C. tortuganus) were reared in the presence of conspecific cocoons, cocoons of the other species, or were kept without cocoons. Groups of older workers (> 20 days), previously exposed to conspecific brood in their natal nest, were familiarized with either conspecific or heterospecific cocoons. After 14 days of exposure, groups were subjected to short-term (5 min) and long-term (10 day) preference tests. Young and older workers retrieved and retained many cocoons of both species, familiar and unfamiliar. However, a pattern of non-exclusive discrimination emerged: (1) Young workers exposed to conspecific cocoons picked up and retrieved conspecifics before unfamiliar heterospecifics, and retained conspecifics longer. (2) Young workers generally required experience with conspecifics to develop this preference, as those exposed to only heterospecifics and those deprived of cocoons were impartial in short-term tests. However, in long-term tests (5–10 days), naive young workers significantly preferred unfamiliar conspecifics. (3) Older workers preferred conspecific cocoons, whether familiar or unfamiliar, in short-term tests, but their tolerance for heterospecifics in the longer term (5 days) could be increased by recent familiarization. In no case did young or older ants significantly prefer familiar heterospecifics to conspecifics. These results confirm a role for early learning in brood recognition by carpenter ants, but suggest that it is less important than in Formica species studied by previous authors.     

Nocturnal orientation in the black carpenter antCamponotus pennsylvanicus (DeGeer) (Hymenoptera: Formicidae)

Summary The black carpenter antCamponotus pennsylvanicus (DeGeer), a predominantly nocturnal Formicine ant, responds to a hierarchy of visual and tactile cues when orienting along odor trails at night. Under illumination from moonlight or artificial light, workers rely upon these beacons to mediate phototactic orientation. In the absence of moonlight or artificial lights, ants were able to orient visually to terrestrial landmarks. In the absence of all landmarks, save for overhanging tree branches, ants could negotiate shortcuts or make directional changes in response to visual landmarks presented within the tree canopy on a moonless night. When experimental manipulations placed the ants in total darkness, they could no longer negotiate shortcuts and would resort to thigmotactic orientation along structural guidelines to reach a food source. The hierachical organization of these diverse cues in a foraging strategy is discussed, as well as their adaptive significance toC. pennsyhanicus.     

Nestmate Recognition in Leptothoracine Ants: Testing Fielde's Progressive Odor Hypothesis

Fielde's progressive odor hypothesis postulates that certain hereditary nestmate recognition odors in adult ants change progressively with age such that workers will often not accept older siblings that differ from them in age by as little as 40–60 days unless they eclosed among siblings of that age class and learned their recognition cues. Fielde's hypothesis was based on extensive but unsystematic experimental studies which fail to provide unequivocal evidence for this phenomenon. The present study tested this hypothesis for three closely-related ant species, Leptothorax ambiguus, L. curvispinosus, and L. longispinosus, by introducing workers between groups with known age structures and a minimum age difference of eight months. The results invalidate Fielde's hypothesis for these ants. However, progressive odors may yet be found in other species and may be associated with age polyethism.     

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.     

Plant‐derived differences in the composition of aphid honeydew and their effects on colonies of aphid‐tending ants

In plant–ant–hemipteran interactions, ants visit plants to consume the honeydew produced by phloem‐feeding hemipterans. If genetically based differences in plant phloem chemistry change the chemical composition of hemipteran honeydew, then the plant's genetic constitution could have indirect effects on ants via the hemipterans. If such effects change ant behavior, they could feed back to affect the plant itself. We compared the chemical composition of honeydews produced by Aphis nerii aphid clones on two milkweed congeners, Asclepias curassavica and Asclepias incarnata, and we measured the responses of experimental Linepithema humile ant colonies to these honeydews. The compositions of secondary metabolites, sugars, and amino acids differed significantly in the honeydews from the two plant species. Ant colonies feeding on honeydew derived from A. incarnata recruited in higher numbers to artificial diet, maintained higher queen and worker dry weight, and sustained marginally more workers than ants feeding on honeydew derived from A. curassavica. Ants feeding on honeydew from A. incarnata were also more exploratory in behavioral assays than ants feeding from A. curassavica. Despite performing better when feeding on the A. incarnata honeydew, ant workers marginally preferred honeydew from A. curassavica to honeydew from A. incarnata when given a choice. Our results demonstrate that plant congeners can exert strong indirect effects on ant colonies by means of plant‐species‐specific differences in aphid honeydew chemistry. Moreover, these effects changed ant behavior and thus could feed back to affect plant performance in the field.     

Sensu stricto individual conditioning,and imitation,in the ant Myrmica sabuleti (Hymenoptera: Formicidae)

Summary

Conditioning of isolated ants was attempted in the ant Myrmica sabuleti. Isolated ants did not die and could be visually as well as olfactorily conditioned. They acquired and kept ? or partly lost ? their visual ? or their olfactory ? conditioned responses as do ants living in a colony. Each individual of an ant colony is thus able to learn and memorize by itself. Any collective conditioning, learning and memory actually reflect the individuals’ performance. Naive workers paired with isolated previously conditioned ants apparently acquired olfactory conditioning in a shorter time than isolated ants, reaching a conditioning score identical to that of isolated ants. But they lost their apparent conditioning as soon as the olfactory cue was removed. Thus, imitation of nestmates exists in ants but does not lead, by itself, to learning sensu stricto since nothing of what has been imitated is retained.     

Visual prey discrimination of queen and worker ants by a generalist lizard

Laboratory experiments were conducted to test the hypothesis that widelyforaging generalist lizard,Eumeces okadae, visually discriminates palatable queen ants from unpalatable worker ants. The workers ofCamponotus japonicus andLasius niger were rejected on sight, while the queens of both species were eaten with little prior chemical examination by tongue flicks or licks. Comparison of the lizards' responses towards the workers and wingless queens of similar size indicated that neither body size nor presence or absence of wings accounted for difference in responses toward the 2 ant castes. The lizards probably discriminated different ant castes by body proportions.     

Getting carbohydrates in ants: damage of the young cereal sprouts by ants of the genus Myrmica Latreille, 1804 (Hymenoptera: Formicidae), to obtain plant sap

Summary

Carbohydrate food is of high importance for survival of ant colonies. Ants are known to use sugary excretions (honeydew) of various insects, nectar of floral and extrafloral nectaries, and even sap of some trees. However, the ability of ants to use sap of herbaceous plants has not been mentioned. This is the first evidence that ants of the genus Myrmica can intentionally ‘cut off’ young cereal sprouts to obtain plant sap. The investigation was carried out in a laboratory in 2018 and 2019 and involved three ant species of the genus Myrmica [12 colonies of M. rubra (Linnaeus, 1758); eight colonies of M. ruginodis Nylander, 1846; and five colonies of M. scabrinodis Nylander, 1846]. First observations were made occasionally in 10 ant colonies during the study of ant–aphid interactions. After three days of carbohydrate starvation, ants were supplied with the plants of wheat infested with aphids of Schizaphis graminum (Rondani, 1852). Within the first day in addition to ordinary trophobiotic relations with aphids, the workers of all the studied colonies demonstrated unexpected behaviour: they ‘cut off’ some sprouts and collected sap of these plants. The experimental investigation in 15 ant colonies of various sizes (about 150, 300 and 500 workers) supplied with the plants infested or non-infested with aphids has shown that getting sap of herbs depends greatly on ant colony needs and available resources. The number of damaged plants was much higher both in the larger colonies of ants and in the absence of aphids. This way of getting carbohydrates allows ants to quickly obtain some extra food needed to maintain colony viability and seems to be one of the mechanisms promoting survival of ants in conditions of acute carbohydrate deficiency. At the same time, ants avoid using plant sap when there are more available alternative carbohydrate resources.     

Polygynous supercolonies of the acacia‐ant Pseudomyrmex peperi,an inferior colony founder

In ant–plant protection mutualisms, plants provide nesting space and nutrition to defending ants. Several plant–ants are polygynous. Possessing more than one queen per colony can reduce nestmate relatedness and consequently the inclusive fitness of workers. Here, we investigated the colony structure of the obligate acacia‐ant Pseudomyrmex peperi, which competes for nesting space with several congeneric and sympatric species. Pseudomyrmex peperi had a lower colony founding success than its congeners and thus, appears to be competitively inferior during the early stages of colony development. Aggression assays showed that P. peperi establishes distinct, but highly polygynous supercolonies, which can inhabit large clusters of host trees. Analysing queens, workers, males and virgin queens from two supercolonies with eight polymorphic microsatellite markers revealed a maximum of three alleles per locus within a colony and, thus, high relatedness among nestmates. Colonies had probably been founded by one singly mated queen and supercolonies resulted from intranidal mating among colony‐derived males and daughter queens. This strategy allows colonies to grow by budding and to occupy individual plant clusters for time spans that are longer than an individual queen’s life. Ancestral states reconstruction indicated that polygyny represents the derived state within obligate acacia‐ants. We suggest that the extreme polygyny of Pseudomyrmex peperi, which is achieved by intranidal mating and thereby maintains high nestmate relatedness, might play an important role for species coexistence in a dynamic and competitive habitat.     

Memory and chemical communication in the orientation of two mass-recruiting ant species

Summary The relative contribution of visual and chemical components in the orientation ofLasius niger andIridomyrmex humilis (Argentine ant) workers during mass recruitment to newly discovered food sources is analyzed over short time intervals. While both species orient in response to the trail pheromone, a large number ofL. niger foragers rapidly switch to a more individual orientation, based on their memory of environmental cues.I. humilis workers, on the other hand, predominantly use collective chemical cues. The effect of the number of reinforcements on visual learning and its interference with chemical communication show that olfactory cues always prevail in the Argentine ant. InL. niger, the proportion of ants orienting to visual cues is independent of the trail concentration. Detailed observations of the trail-laying behavior of individually marked foragers show that nearly all theI. humilis workers initially lay a trail, whereas only half theL. niger foragers do so. This proportion decreases considerably with the number of trips performed byL. niger workers, while remaining constant for the Argentine ants. These results are interpreted with respect to the species' behavioral ecology.     

Ant-seed interactions: combined effects of ant and plant species on seed removal patterns

Seed dispersal by ants (i.e. myrmecochory) is usually considered as a mutualism: ants feed on nutritive bodies, called elaiosomes, before rejecting and dispersing seeds in their nest surroundings. While mechanisms of plant dispersal in the field are well documented, the behaviour of the ant partner was rarely investigated in details. Here, we compared in laboratory conditions the foraging behaviour of two ant species, the omnivorous Lasius niger and the insectivorous Myrmica rubra to which seeds of two European myrmecochorous plants (Chelidonium majus and Viola odorata) were given. Ant colonies were simultaneously presented three types of items: entire seeds with elaiosome (SE), seeds without elaiosome (S) and detached elaiosomes (E). The presence of elaiosomes on seeds did not attract workers from a distance since ants first contact equally each type of items. Although ants are mass-recruiting species, we never observed any recruitment nor trail-laying behaviour towards seeds. For ants having contacted seed items, their antennation, manipulation and seed retrieval behaviour strongly varied depending on the species of each partner. Antennation behaviour, followed by a loss of contact, was the most frequent ant-seed interaction and can be considered as a “hesitation” clue. For both plant species, insectivorous Myrmica ants removed items in larger number and at higher speed than Lasius. This fits with the hypothesis of a convergence between odours of elaiosomes and insect preys. For both ant species, the small Chelidonium seeds were retrieved in higher proportion than Viola ones, confirming the hypothesis that ants prefer the higher elaiosome/diaspore-ratio. Thus, in these crossed experiments, the ant-plant pair Myrmica/Chelidonium was the most effective as ants removed quickly almost all items after a few antennations. The presence of an elaiosome body increased the seed removal by ants excepting for Myrmica which retrieved all Chelidonium seeds, even those deprived of their elaiosome. After 24 h, all the retrieved seeds were rejected out of the nest to the refuse piles. In at least half of these rejected items, the elaiosome was discarded by ants. Species-specific patterns and behavioural differences in the dynamics of myrmecochory are discussed at the light of ant ecology. Received 10 September 2007; revised 5 February 2008; accepted 5 March 2008.     

Meat ants as dominant members of Australian ant communities: an experimental test of their influence on the foraging success and forager abundance of other species

Meat ants (Iridomyrmex purpureus and allies) are perceived to be dominant members of Australian ant communities because of their great abundance, high rates of activity, and extreme aggressiveness. Here we describe the first experimental test of their influence on other ant species, and one of the first experimental studies of the influence of a dominant species on any diverse ant community. The study was conducted at a 0.4 ha savanna woodland site in the seasonal tropics of northern Australia, where the northern meat ant (I. sanguineus) represented 41% of pitfall catches and 73% of all ants at tuna baits, despite a total of 74 species being recorded. Meat ants were fenced out of experimental plots in order to test their influence on the foraging success of other species, as measured by access to tuna baits. The numbers of all other ants and ant species at baits in exclusion plots were approximately double those in controls (controlling for both the fences and for meat ant abundance), and returned rapidly to control levels when fences were removed after 7 weeks. Individual species differend markedly in their response to the fencing treatment, with species of Camponotus and Monomorium showing the strongest responses. Fencing had no effect on pitfall catches of species other than the meat ant, indicating that the effect of meat ants at baits was directly due to interference with foraging workers, and not regulation of general forager abundance. Such interference by meat ants has important implications for the sizes and densities of colonies of other ant species, and ultimately on overall ant community structure.     

Imprinting Effects on Host-selection Behavior of Slave-raiding Chalepoxenus muellerianus (Finzi) Workers (Hymenoptera: Formicidae)

Early experience is known to influence brood care behavior and to induce nesting preferences of several ant species. Scanty information is available with respect to imprinting effects on host selection of dulotic ants. In this paper we investigate host choice behavior of workers of the slave-making ant, Chalepoxenus muellerianus. As in a corresponding paper (Schumann & Buschinger 1994) on host specificity of C. muellerianus young queens during colony foundation, we demonstrate that a blend of innate factors and effects of early experience also influences host species selection of slave-raiding workers.     

A single lycaenid caterpillar gets an ant‐constructed shelter and uninterrupted ant attendance

Ant‐lycaenid associations range from mutualism to parasitism and the caterpillars of some species of lycaenids are reported to enter ant nests for shelter, diapause, or pupation. The present study aimed to examine the nature of the association between Euchrysops cnejus (Fabricius) (Lepidoptera: Lycaenidae) and Camponotus compressus (Fabricius) (Hymenoptera: Formicidae) worker ants on the extrafloral nectary‐bearing cowpea plant, Vigna unguiculata (L.) Walp. (Fabaceae). The abundance patterns of the ants and the lycaenid caterpillars together with the spatial patrolling patterns of the ants on the plants revealed that ant abundance increased with the occurrence of the lycaenid caterpillars and the ants preferred the lycaenids over the extrafloral nectar. Camponotus compressus worker ants constructed a shelter at the cowpea plant base after interaction with one or more lycaenid caterpillar(s) and tended the caterpillars and pupae till the emergence of the butterfly. The ant‐constructed shelters (ACSs) inhabited by the minor caste workers (13 ± 1.3 ants per ACS), were utilized by the caterpillars to undergo pupation. The ants confined their activities predominantly to tending the pod‐feeding caterpillars and the solitary pupa within each ACS. It appears that the behavior of the tending worker ants is modulated by the lycaenid vulnerable stages.     

Phorid parasitoids affect foraging activity of Solenopsis richteri under different availability of food in Argentina

1. In Argentina, six species of Pseudacteon parasitoids (Phoridae) attack Solenopsis richteri, one of the two species of South American fire ant that are exotic pests in North America. 2. The presence of these Pseudacteon species significantly reduces the number of ants at food resources in the field, as well as foraging activity generally. 3. Some Pseudacteon not only attack ants walking on trails or at feeding sites, but also at mound entrances, inhibiting workers from leaving to forage. 4. The average size of foraging ants (which prescribes their suitability as hosts) decreased in the presence of phorids. 5. The number of attacking phorids was correlated positively with the number of ants walking towards the food on the trail before the attack. 6. Solenopsis richteri workers responded to manipulations of food size and presence or absence of parasitoids in a risk-adjusting way, i.e. although more foragers were recruited to larger food items, attacking phorids reduced ant foraging activity by the same factor regardless of the size of the food offered. 7. The data suggest that S. richteri colonies juggle the needs to harvest food efficiently, reduce competition, and avoid excess risks from parasitoids in complex ways.     

Thorn‐dwelling ants provide antiherbivore defence for camelthorn trees,Vachellia erioloba,in Namibia

Ants are widely employed by plants as an antiherbivore defence. A single host plant can associate with multiple, symbiotic ant species, although usually only a single ant species at a time. Different plant‐ant species may vary in the degree to which they defend their host plant. In Kenya, ant–acacia interactions are well studied, but less is known about systems elsewhere in Africa. A southern African species, Vachellia erioloba, is occupied by thorn‐dwelling ants from three different genera. Unusually, multiple colonies of all these ants simultaneously and stably inhabit trees. We investigated if the ants on V. erioloba (i) deter insect herbivores; (ii) differ in their effectiveness depending on the identity of the herbivore; and (iii) protect the tree against an important herbivore, the larvae of the lepidopteran Gonometa postica. We show that experimental exclusion of ants leads to greater levels of herbivory on trees. The ants inhabiting V. erioloba are an effective deterrent against hemipteran and coleopteran, but not lepidopteran herbivores. Defensive services do not vary among ant species, but only Crematogaster ants exhibit aggression towards G. postica. This highlights the potential of the V. erioloba–ant mutualism for studying ant–plant interactions that involve multiple, simultaneously resident thorn‐dwelling ant species.