Burning bridges: priority effects and the persistence of a competitively subordinate acacia-ant in Laikipia, Kenya

Theoretical models suggest that subordinate competitors may rely on strong colonization ability and/or high persistence (e.g., the ability to resist invasion) as a strategy to coexist with competitively dominant species. While strong colonization ability among subordinate competitors has been widely documented, we know less about the role of persistence in facilitating species coexistence. In upland East Africa, four species of acacia-ants (Crematogaster sjostedti, C. mimosae, C. nigriceps, Tetraponera penzigi) compete for possession of Acacia drepanolobium host trees. Despite a strong dominance hierarchy, the four acacia-ant species coexist at fine spatial scales. Here we present evidence that T. penzigi, the least aggressive competitor, modifies host trees in two ways that reduce the probability of aggressive takeover by neighboring colonies. First, T. penzigi workers destroy virtually all leaf nectaries on their host trees. Second, T. penzigi workers create and maintain entryways into their swollen thorn domiciles that are too small to allow entry by their Crematogaster competitors. In a 2Ƕ factorial experiment, we manipulated nectar availability and swollen-thorn entryway size to determine the influence of these factors on the probability of aggressive displacement by a dominant competitor (C. mimosae) in staged conflicts. Addition of artificial nectaries and enlargement of swollen-thorn entryways on T. penzigi-occupied trees increased the probability of aggressive displacement of T. penzigi by C. mimosae from swollen thorns 14-fold and 8-fold, respectively. Further, empty saplings with nectaries destroyed by T. penzigi workers were colonized by half as many C. mimosae workers as saplings where nectaries were left intact. Our results demonstrate that T. penzigi's unusual strategy of nectary destruction and the maintenance of small entryways in swollen thorns produce priority effects, effectively reducing the probability that T. penzigi colonies will be displaced from host trees by more dominant competitors.     

Development rate and brood production in haplo- and pleometrotic colonies of <Emphasis Type="Italic">Oecophylla smaragdina</Emphasis>

Pleometrosis (colony founding by multiple queens) may improve life history characteristics that are important for early colony survival. When queens unite their initial brood, the number of workers present when incipient colonies open may be higher than for single queen colonies. Further, the time until the first worker emerges may shorten. For territorial species and species that rob brood from neighbouring colonies, a faster production of more workers may improve the chance of surviving intraspecific competition. In this study, the time from the nuptial flight to the emergence of the first worker in incipient Oecophylla smaragdina Fabr. colonies founded by 1–5 queens was compared and the production of brood during the first 68 days after the nuptial flight was assessed. Compared to haplometrotic colonies, pleometrotic colonies produced 3.2 times more workers, their first worker emerged on average 4.3 days (8%) earlier and the queen’s per capita egg production almost doubled. Further, colony production was positively, correlated with the number of founding queens and time to worker emergence was negatively correlated. These results indicate that pleometrotic O. smaragdina colo-nies are competitively superior to haplometrotic colonies as they produce more workers faster and shorten the claustral phase, leading to increased queen fecundity.     

The reproductive phenology of an Amazonian ant species reflects the seasonal availability of its nest sites

In saturated tropical ant assemblages, reproductive success depends on queens locating and competing for scarce nest sites. Little is known about how this process shapes the life histories of tropical ants. Here I investigate the relationship between nest site availability and an important life history trait, reproductive phenology, in the common Amazonian ant species Allomerus octoarticulatus. A. octoarticulatus is a plant-ant that nests in the hollow, swollen stem domatia on Cordia nodosa. I provide evidence that nest sites are limiting for A. octoarticulatus. Most queens produced by A. octoarticulatus colonies died before locating suitable host plants, and most queens that located hosts died before founding colonies, probably from intraspecific competition among queens for control of host plants. I further show that the reproductive phenology of A. octoarticulatus closely matches the seasonal availability of its nest sites, domatia-bearing C. nodosa saplings. Both the production and flight of A. octoarticulatus reproductives, and the number of C. nodosa saplings available for colonization by ants, peaked from March to May. There was correlative evidence that A. octoarticulatus colonies use temperature as a cue to synchronize their reproduction to the availability of C. nodosa saplings: both the production of reproductives by ant colonies and the number of C. nodosa saplings available for colonization were correlated with temperature, and not with rainfall. All of these results suggest that nest site limitation constrains the reproductive phenology of A. octoarticulatus.     

A shift in colony founding behaviour in the obligate plant-ant Crematogaster (Decacrema) morphospecies 2

Summary. A shift in colony founding behaviour from single queen (haplometrosis) to multiple queens (pleometrosis) was observed locally in the obligate plant-ant Crematogaster (Decacrema) morphospecies 2, which is associated with Macaranga trees in Borneo. In addition, about a quarter of all mature colonies (27 of 95 trees examined) were found to be multiple queen colonies. They arose either directly from pleometrotic founding colonies or secondarily by adoption of additional queens. Using microsatellite markers, we showed that queens in colonies founded through pleometrosis are unrelated and each queen participates in producing worker offspring, albeit with significant skew in a third of the colonies. In mature polygynous colonies, all resident queens contributed to the production of workers and sexual offspring. Relatedness of queens in mature polygynous colonies was not significantly higher than in foundress associations. We hypothesize that increased nest site limitation in this specific interaction trigger the observed shift in colony founding behaviour. Crematogaster msp. 2 inhabits the light demanding pioneer plant species Macaranga pearsonii that is typical for early successional stages of secondary forests. Thus suitable host-plants for colonisation are abundant for only a short time in highly disturbed sites and become increasingly sparse when secondary forest matures.Received 6 September 2004; revised 29 November; accepted 10 December 2004.     

Polygyny does not explain the superior competitive ability of dominant ant associates in the African ant‐plant,Acacia (Vachellia) drepanolobium

The Acacia drepanolobium (also known as Vachellia drepanolobium) ant‐plant symbiosis is considered a classic case of species coexistence, in which four species of tree‐defending ants compete for nesting space in a single host tree species. Coexistence in this system has been explained by trade‐offs in the ability of the ant associates to compete with each other for occupied trees versus the ability to colonize unoccupied trees. We seek to understand the proximal reasons for how and why the ant species vary in competitive or colonizing abilities, which are largely unknown. In this study, we use RADseq‐derived SNPs to identify relatedness of workers in colonies to test the hypothesis that competitively dominant ants reach large colony sizes due to polygyny, that is, the presence of multiple egg‐laying queens in a single colony. We find that variation in polygyny is not associated with competitive ability; in fact, the most dominant species, unexpectedly, showed little evidence of polygyny. We also use these markers to investigate variation in mating behavior among the ant species and find that different species vary in the number of males fathering the offspring of each colony. Finally, we show that the nature of polygyny varies between the two commonly polygynous species, Crematogaster mimosae and Tetraponera penzigi: in C. mimosae, queens in the same colony are often related, while this is not the case for T. penzigi. These results shed light on factors influencing the evolution of species coexistence in an ant‐plant mutualism, as well as demonstrating the effectiveness of RADseq‐derived SNPs for parentage analysis.     

Patterns of colonization and incipient nest survival inAcromyrmex niger andAcromyrmex balzani (Hymenoptera: Formicidae)

Summary Patterns of colonization by queens and incipient nest survival of the leaf-cutting antsAcromyrmex niger andAcromyrmex balzani were studied by means of belt transects and individually marked incipient nests. No relation was found between colony density and the number of colonization attempts. Both species are not claustral, and high rates of queen mortality were attributed to conspecific executions and predation. Of other discernable mortality factors, failure of fungal garden establishment was the most important. Only 34 of 296A. balzani and 13 of 154A. niger marked colonies were alive at the end of one year. These figures are higher than those reported for species ofAtta. These results are contrasted with those of claustral-foundingAtta species. Small colonies are occasionally raided by larger colonies which robbed brood.     

Polygyny in the nest-site limited acacia-ant Crematogaster mimosae

Polygyny is common in social insects despite inevitable decreases in nestmate relatedness and reductions to the inclusive fitness returns for cooperating non-reproductive individuals. We studied the prevalence and mode of polygyny in the African acacia-ant Crematogaster mimosae. These ants compete intensively with neighboring colonies of conspecifics and with three sympatric ant species for resources associated with the whistling-thorn acacias in which they all obligately nest. We used the genotypes of alate males at ten microsatellite loci to reconstruct queen genotypes and found that C. mimosae colonies are frequently secondarily polygynous, in that they include multiple closely related (and sometimes full-sib) queens, and (more rarely) unrelated queens. We also found that individual queens in both monogynous and polygynous colonies had mated with multiple males, making C. mimosae an interesting example of simultaneous polygyny and polyandry. The presence of polygyny in C. mimosae and the intense competition for nest-sites between C. mimosae and its conspecifics support the association between nest-site limitation and polygyny. Polygyny may allow for increased worker populations and a competitive advantage, as inter-colony conflicts are typically won by the colony with the larger number of workers.     

An experimental analysis of pleometrotic advantage in the desert seed-harvester antMessor pergandei (Hymenoptera; Formicidae)

Summary Starting colonies of the desert seed-harvester antMessor pergandei are clumped in the field and face severe intraspecific competition through brood raiding. Single foundress laboratory colonies ofM. pergandei are more likely to succeed at brood raiding with conspecific colonies if they are given additional workers and mature pupae several days prior to brood raiding. Per foundress fecundity remains constant across laboratory starting colonies established with 1, 3 and 5 foundresses. These results suggest that the selective advantage of cooperative colony foundation (pleometrosis) in this and similar species may derive directly from the ability of multiple foundresses to produce a larger brood raiding force.     

Cooperative foundation of colonies by unrelated foundresses in the antPolyrhachis moesta

Summary Field studies ofPolyrhachis moesta revealed that colonies were facultatively founded by more than one queen, even though each established colony was inhabited by only one queen with her workers. In pleometrotic colonies, it was found that cofoundresses frequently exhibited cooperative behaviors, including trophallaxis. The results of DNA fingerprinting showed that multiple foundresses were not genetically to one another. It seems that they aggregate randomly and independently of their relatedness. A discussion of the reasons why foundresses ofP. moesta do not aggregate preferentially with genetically related females.     

Mating flight, metrosis, and semi-claustrality in the seed-harvester ant Pogonomyrmex salinus (Hymenoptera, Formicidae)

Here we examine dispersal, metrosis, and claustrality in the seed-harvester ant Pogonomyrmex salinus at an unusually large mating aggregation. We found that mode of queen dispersal from the mating aggregation is not a function of queen mass and that wing damage among queens was relatively rare. P. salinus is haplometrotic in the field and foundress queens placed together in forced associations eventually fight to the death. While queens of Pogonomyrmex salinus can survive claustrally, producing a single minim from their body reserves in the laboratory, fed queens produce up to four significantly larger minims along with concurrent larvae and pupae during the same period. Since queens forage in the field, we interpret claustrality as a secondary reserve strategy when foraging fails, and suggest that foraging is obligate for P. salinus queens in an overdispersed and temperate environment. Thus, nest founding strategies employed by P. salinus may be environmentally determined and represent a continuum between fully claustral and obligate foraging. We discuss our results with reference to theories of pleometrosis and claustral colony founding. Received 12 November 2004; revised 12 April 2004; accepted 29 July 2005.     

Infestation by a Common Parasite is Correlated with Ant Symbiont Identity in a Plant‐Ant Mutualism

In East Africa, up to four symbiotic ant species associate with the obligate myrmecophyte Acacia drepanolobium. These ant species differ in the extent to which they defend their host trees from both vertebrate and invertebrate herbivores, but other potential roles of ants in tree defense have not been studied. We investigate the distribution of a new species of parasitic midge targeting A. drepanolobium in a region where A. drepanolobium is inhabited almost exclusively by two ant species—Crematogaster nigriceps and C. mimosae. We find that the frequency of infestation correlates strongly with the identity of the ant occupant: trees inhabited by C. nigriceps are significantly less likely to be infested with parasitic midges. Although the two ant species responded similarly to simulated large herbivore disturbances, trees inhabited by C. nigriceps also had a lower invertebrate load than trees inhabited by C. mimosae. We suggest that differences in defensive behavior towards invertebrates could be one explanation of the observed differences in infestation of A. drepanolobium by parasitic midges.     

Number of queens in founding associations of the ponerine ant Pachycondyla villosa

In the ant Pachycondyla villosa, new colonies are usually started cooperatively by two or more young queens who establish a dominance order with a division of labour. Co-founding can lead to primary polygyny, where queens stay together after workers have emerged. Here we show that two queens associations are the most common (47%) and also the most stable in the field. When offered additional nest sites in the laboratory, two-queen associations did not split, whereas larger associations did so. We also show that solitary foundresses always accepted experimentally introduced alien queens, while these were attacked and sometimes killed in queen associations. The removal of dominant alpha queens from three-queen associations resulted in beta queens obtaining the dominant role and sometimes the destruction of the existing eggs. It appears that two queens suffice for a successful association and that pleometrosis is favoured by ecological constrains, such as nest-site limitation. Received 28 February 2005; revised 20 April 2005; accepted 22 April 2005.     

Energetics of newly-mated queens and colony founding in the fungus-gardening ants Cyphomyrmex rimosus and Trachymyrmex septentrionalis (Hymenoptera: Formicidae)

Abstract.  The energetics of colony founding is investigated in the fungus gardening ants (Attini) Trachymyrmex septentrionalis and Cyphomyrmex rimosus . Similar to most ants, inseminated queens of these two species found nests independently unaccompanied by workers (haplometrosis). Whereas most ant founding queens seal themselves in a chamber and do not feed when producing a brood entirely from metabolic stores (claustral founding), the majority of fungus gardening ants must forage during the founding phase (semiclaustral founding). Laboratory-reared T. septentrionalis individuals comprise 84 dealate females collected after mating flights in June 2004. Twenty are immediately killed to obtain values for queen traits and another 20 after worker emergence for queen, fungus garden and worker traits. Cyphomyrmex rimosus comprise 22 dealate females collected in June 2005; ten of which are immediately killed and similarly prepared. Newly-mated T. septentrionalis queens have 25% of their dry weight as fat; whereas newly-mated C. rimosus queens contain 11% fat. These amounts are 50–75% less than most independently founding ant species. Trachymyrmex septentrionalis queens lose merely 5% of their energetic content during colony founding, whereas the total energetic content of their brood is more than three-fold the amount lost by the queen. Incipient T. septentrionalis colonies produce approximately half as much ant biomass per gram of fungus garden as do mature colonies. Similar to most ants, T. septentrionalis produces minim workers that are approximately 40% lighter than workers from mature colonies. Regardless of their size, T. septentrionalis workers contain much lower fat than do workers of claustral species. These data indicate that fungus gardening is adaptive because colonies can produce much cheaper offspring, making colony investment much lower.     

Nutritional benefits of crematogaster mimosae ants and acacia drepanolobium gum for patas monkeys and vervets in laikipia,Kenya

Patas monkeys (Erythrocebus patas) are midsized primates that feed extensively on the gum of Acacia drepanolobium and the ants are housed in swollen thorns of this Acacia. Their diet resembles that expected more of smaller bodied primates. Patas monkeys are also more like smaller bodied primates in reproducing at high rates. We sought to better understand the convergence of patas monkeys with smaller bodied primates by comparing their feeding behavior on ants and gum with that of closely related, sympatric vervets (Chlorocebus pygerythrus), and analyzing the nutrient content of the gum of A. drepanolobium and of Crematogaster mimosae, the most common ant species eaten by patas monkeys in Laikipia, Kenya. All occurrences of feeding and moving during focal animal sampling revealed that 1) patas monkeys seek A. drepanolobium gum but vervets avoid it; 2) both species open swollen thorns most often in the morning when antsare less active; 3) patas monkeys continually feed onswollen thorns and gum while moving quickly throughout the day, whereas vervets reduce their consumption of these items and their travel rate at mid‐day, and; 4) vervets eat young swollen thorns at a higher rate than patas monkeys. Patas monkeys are able to spend little time acquiring substantial amounts of energy, protein, and minerals from A. drepanolobium gum and C. mimosae ants each day. These findings, when coupled with evidence of causes of infant and adult female mortality, suggest that reproductive success of female patas monkeys is more immediately affected by illness, disease, interactions between adults and infants, and access to water than by food. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.     

The cost of being queen: Investment across Pogonomyrmex harvester ant gynes that differ in degree of claustrality

The role of the ant colony largely consists of non-reproductive tasks, such as foraging, tending brood, and defense. However, workers are vitally linked to reproduction through their provisioning of sexual offspring, which are produced annually to mate and initiate new colonies. Gynes (future queens) have size-associated variation in colony founding strategy (claustrality), with each strategy requiring different energetic investments from their natal colony. We compared the per capita production cost required for semi-claustral, facultative, and claustral gynes across four species of Pogonomyrmex harvester ants. We found that the claustral founding strategy is markedly expensive, costing approximately 70% more energy than that of the semi-claustral strategy. Relative to males, claustral gynes also had the largest differential investment and smallest size variation. We applied these investment costs to a model by Brown and Bonhoeffer (2003) that predicts founding strategy based on investment cost and foraging survivorship. The model predicts that non-claustral foundresses must survive the foraging period with a probability of 30–36% in order for a foraging strategy to be selectively favored. These results highlight the importance of incorporating resource investment at the colony level when investigating the evolution of colony founding strategies in ants.     

Scaling of body weight and fat content in fungus-gardening ant queens: does this explain why leaf-cutting ants found claustrally?

Offspring traits are among the most important life history traits, yet we lack an adequate understanding of their role in social insect life history evolution. Colony founding in the fungus-gardening ants (Tribe Attini) is different from most other ant species because the queens forage during the founding phase. Queens of the most derived genus, Atta, are the only attines that exhibit the more typical claustral founding, where the queens seal themselves in a below-ground chamber and produce their first generation of workers with only body fat reserves. Here I report the dry weights, fat content and energetic value of newly mated queens of ten attine species. Published phylogenies were used to make inferences on the evolutionary transitions in this clade. It appears that the evolution of fungus-gardening was associated with the manufacture of smaller, leaner queens as basal taxa are characterized by small bodies that contain relatively less fat than derived taxa. Moreover, there appears to be an allometric function between fat, energetic content and dry weight, which means that for fatter and claustral queens to develop, they also must become larger.     

Modes of colony foundation by females of different morphotypes in the paper wasps (Hymenoptera,Vespidae, <Emphasis Type="Italic">Polistes</Emphasis> Latr.)

Paper wasps of the genus Polistes exhibit wide variability both of the color and size of foundresses, and the mode of colony foundation. The colony foundation and social hierarchy were studied in populations of two Palaearctic species, Polistes dominulus (Christ) and P. nimphus (Christ) (April–May 2006, the Crimean Peninsula, Ukraine), and one Neotropical species, P. lanio (F.) (October 2004–January 2005, Trinidad Island). It was shown that foundresses of various color and size morphs tended to found nests in a peculiar way: singly (haplometrosis), by groups (pleometrosis), or occupying several nests simultaneously (polycaly). The relationship between coloration, size, and mode of colony foundation was species-specific. P. dominulus foundresses with a darker clypeus and a paler mesonotum preferred to found nests alone and had a dominant position in pleometrosis. The haplometrotic P. nimphus foundresses more often displayed darker variants of the clypeus and paler variants of the mesonotum than did foundresses from pleometrotic colonies. P. lanio foundresses from pleometrotic colonies differed from those from polycalic colonies in the coloration variability of the scutum, propodeum, and the 2nd metasomal tergite. The dominant and subordinate P. nimphus and P. lanio foundresses differed in the size of head and wings. The possible significance of the subdivision of foundresses into the “generalists” and “specialists” for the variability structuring in the population is considered.     

Dispersal Polymorphisms in Invasive Fire Ants

In the Found or Fly (FoF) hypothesis ant queens experience reproduction-dispersal tradeoffs such that queens with heavier abdomens are better at founding colonies but are worse flyers. We tested predictions of FoF in two globally invasive fire ants, Solenopsis geminata (Fabricius, 1804) and S. invicta (Buren, 1972). Colonies of these species may produce two different monogyne queen types—claustral queens with heavy abdomens that found colonies independently, and parasitic queens with small abdomens that enter conspecific nests. Claustral and parasitic queens were similarly sized, but the abdomens of claustral queens weighed twice as much as those of their parasitic counterparts. Their heavier abdomens adversely impacted morphological predictors of flight ability, resulting in 32–38% lower flight muscle ratios, 55–63% higher wing loading, and 32–33% higher abdomen drag. In lab experiments maximum flight durations in claustral S. invicta queens decreased by about 18 minutes for every milligram of abdomen mass. Combining our results into a simple fitness tradeoff model, we calculated that an average parasitic S. invicta queen could produce only 1/3 as many worker offspring as a claustral queen, but could fly 4 times as long and have a 17- to 36-fold larger potential colonization area. Investigations of dispersal polymorphisms and their associated tradeoffs promises to shed light on range expansions in invasive species, the evolution of alternative reproductive strategies, and the selective forces driving the recurrent evolution of parasitism in ants.     

Preparation for disturbance-induced absconding of Cape honeybee colonies (Apis mellifera capensis Esch.)

African honeybees, Apis mellifera, are characterised by frequent disturbance-induced absconding. However, the effectiveness in preparation before such disturbance-induced absconding has not been rigorously quantified yet. We investigated the effectiveness of preparation for disturbance-induced absconding by evaluating colony phenotypes prior to and after absconding in ten colonies of the Cape honeybee, A. m. capensis. Seven non-absconding colonies at the same apiary were used as controls. While seven absconded colonies left neither stores nor brood behind, three colonies abandoned only a small area of honey, pollen, open or capped brood. At the end of the observations, the control colonies still had pollen and honey stores and brood. The mean reduction rate between a major disturbance and the absconding event was 0.052 ± 0.018 cm² stores and open brood per worker per day. Our results demonstrate that disturbance-induced absconding can also occur with preparation, if the disturbance is not highly destructive and enough time for preparation is available. We conclude that Cape honeybee colonies can show a considerable high effectiveness in their preparation before disturbance-induced absconding, which limits the loss of colony resources. In light of the general high mobility of African honeybee colonies such an efficient behaviour is probably adaptive. Received 22 December 2004; revised 3 June 2005; accepted 13 June 2005.     

Hierarchical Perception of Stimuli During Case Construction in the Bagworm Moth Eumeta crameri (Lepidoptera: Psychidae)

We studied case construction behavior of bagworm moth, Eumeta crameri (Lepidoptera: Psychidac) in the host plant, Acacia nilotica, at two sites. At the time of the study, Site A had 1-year-old A. nilotica saplings only, whereas Site B had full grown trees. The larva of bagworm moth used either thorns or cut-twigs or both as the materials for building its case. It renovated its case three times, and during each instance it increased the volume of its case by replacing older thorns or cut-twigs by newer and longer ones. However, it exhibited a spectacular predilection for thorns, irrespective of the sites, during the first-instar stage of its development. Thereafter, at Site A it used exclusively cut-twigs, whereas at Site B it preferably used thorns provided they were available in the range of the required length and within the threshold distance. It appears that the bagworm has an ability to process thorn length and distance signals hierarchically to fulfill its priority, that is, case building. Thus it optimizes time and energy expenditures during the period of its larval growth leading to pupation by toggling its preference between thorns and cut-twigs.