Object Depots in the Genus <Emphasis Type="Italic">Pogonomyrmex</Emphasis>: Exploring the “Who,” What,When, and Where

Harvester ants of the genus Pogonomyrmex collect and deposit many items on top of their nests. The depots of P. badius consist mostly of small charcoal fragments, while those of other species are primarily pebbles. Mature colonies can have hundreds of thousands of objects in their depot. In P. badius, the distributions of midden and charcoal about the mound are not completely overlapping, but are positively correlated in areas of overlap. Charcoal depots are isometric with colony size, but the amount of charcoal per colony size varies with season and site. The absence of the depot stimulates collection of nonfood objects. Recruitment to objects only occurs in the presence of food, and foragers choose objects based on size, but not color. An overview of current knowledge concerning depots in the genus indicates that depot formation is likely the ancestral state in the North American species. Furthermore, it is likely that selection is operating indirectly on these depots through selection on many dependent tasks.     

The short-term regulation of foraging in harvester ants

In the seed-eating ant Pogonomyrmex barbatus, the return ofsuccessful foragers stimulates inactive foragers to leave thenest. The rate at which successful foragers return to the nestdepends on food availability; the more food available, the morequickly foragers will find it and bring it back. Field experimentsexamined how quickly a colony can adjust to a decline in therate of forager return, and thus to a decline in food availability,by slowing down foraging activity. In response to a brief, 3-to 5-min reduction in the forager return rate, foraging activityusually decreased within 2–3 min and then recovered within5 min. This indicates that whether an inactive forager leavesthe nest on its next trip depends on its very recent experienceof the rate of forager return. On some days, colonies respondedmore to a change in forager return rate. The rapid colony responseto fluctuations in forager return rate, enabling colonies toact as risk-averse foragers, may arise from the limited intervalover which an ant can track its encounters with returning foragers.     

Polyandry in two South American harvester ants

Although monandry (single mating) is the ancestral state in social hymenopteran insects, effective mating frequencies greater than 2 have been confirmed for a fair amount of ant species: Cataglyphis cursor, the leaf-cutters of the genera Atta and Acromyrmex, army ants of the genera Eciton, Dorylus, Aenictus and Neivamyrmex, and some North American seed harvester species of the genus Pogonomyrmex. This last genus spreads throughout open arid habitats from Patagonia to southwestern Canada. Whereas some North American Pogonomyrmex species are thoroughly studied, we know much less about these ants in South America. The objective of this study was to estimate the effective mating frequency of Pogonomyrmex inermis and P. pronotalis, two Pogonomyrmex sensu stricto species from the central Monte desert of Argentina. A total of 477 P. pronotalis workers from 24 colonies and 402 P. inermis workers from 20 colonies were analyzed using six and four highly polymorphic microsatellites, respectively. The multilocus analysis revealed that all colonies were monogynous and all queens multiply-mated. The effective mating frequency was 8.75 and 6.52 for queens of P. pronotalis and P. inermis, respectively; those values increased up to 15.66 and 9.78, respectively, when corrected for sampling errors. This is the first demonstration that queens in at least some members of the South American Pogonomyrmex sensu stricto are strictly polyandrous, with mating numbers per queen at least as high as those previously found for North American species. We suggest that multiple mating probably arose early in the evolution of the genus Pogonomyrmex and may be the basis of its ecological success and wide distribution. Received 11 October 2006; revised 10 August 2007 and 19 November 2007; accepted 21 November 2007.     

The effects of colony characteristics on life span and foraging behavior of individual wasps (Polybia occidentalis,Hymenoptera: Vespidae)

Summary We studied the effects of intrinsic colony characteristics and an imposed contingency on the life span and behavior of foragers in the swarm-founding social waspPolybia occidentalis. Data were collected on marked, known-age workers introduced into four observation colonies.To test the hypothesis that colony demographic features affect worker life span, we examined the relationships of colony age and size with worker life span using survivorship analysis. Colony age and size had positive relationships with life span; marked workers from two larger, older colonies had longer life spans (¯X = 24.7 days) than those from two smaller, younger colonies (¯X = 20.1 days).We quantified the effects of experimentally imposed nest damage on forager behavior, to determine which of three predicted behavioral responses by foragers to this contingency (increased probability of foraging for building material, increased rate of foraging, or decrease in age of onset of foraging) would be employed. Increasing the colony level of need for materials used in nest construction (wood pulp and water) by damaging the nests of two colonies did not cause an increase in either the proportion of marked workers that gathered nest materials or in foraging rates of marked individuals, when compared with introduced workers in two simultaneously observed control colonies. Instead, nest damage caused a decrease in the age at which marked workers first foraged for pulp and water. The response to an increase in the need for building materials was an acceleration of behavioral development in some workers.     

Evolution of Host Search Strategies in Entomopathogenic Nematodes

There is interspecific variation in infective juvenile behavior within the entomopathogenic nematode genus Steinernema. This variation is consistent with use of different foraging strategies along a continuum between ambush and cruise foraging. To address questions about the evolution of foraging strategy, behavioral and morphological characters were mapped onto a phylogeny of Steinernema. Three species, all in the same clade, were classified as ambushers based on standing bout duration and host-finding ability. One clade of six species were all cruisers based on both host-finding and lack of standing behavior. All species in the ambusher clade had a high rate of jumping, all species in the cruiser clade had no jumping, and most intermediate foragers exhibited some level of jumping. Response to volatile and contact host cues was variable, even within a foraging strategy. Infective juveniles in the ambusher clade were all in the smallest size category, species in the cruiser clade were in the largest size categories, and intermediate foragers tended to be more intermediate in size. We hypothesize that the ancestral Steinernema species was an intermediate forager and that ambush and cruise foraging both evolved at least once in the genus.     

Quantifying the effect of colony size and food distribution on harvester ant foraging

Desert seed-harvester ants, genus Pogonomyrmex, are central place foragers that search for resources collectively. We quantify how seed harvesters exploit the spatial distribution of seeds to improve their rate of seed collection. We find that foraging rates are significantly influenced by the clumpiness of experimental seed baits. Colonies collected seeds from larger piles faster than randomly distributed seeds. We developed a method to compare foraging rates on clumped versus random seeds across three Pogonomyrmex species that differ substantially in forager population size. The increase in foraging rate when food was clumped in larger piles was indistinguishable across the three species, suggesting that species with larger colonies are no better than species with smaller colonies at collecting clumped seeds. These findings contradict the theoretical expectation that larger groups are more efficient at exploiting clumped resources, thus contributing to our understanding of the importance of the spatial distribution of food sources and colony size for communication and organization in social insects.     

Effect of leaf toughness on the susceptibility of the leaf-cutting ant Atta sexdens to attacks of a phorid parasitoid

Summary: Because the size of Atta spp. along foraging trails is partly determined by the characteristics of the plants harvested, and considering that parasitic phorid flies are attracted mostly to large individuals, we hypothesized that plant toughness affects the susceptibility of Atta spp. to these parasitoids. To test this hypothesis, we evaluated parasitism rates of the phorid Neodohrniphora sp. and its effect on Atta sexdens (L.) foragers in a laboratory colony. We manipulated forager size by alternating tough (Anthocephalus chinensis, Rubiaceae) and tender (Rosa chinensis, Rosaceae) plants given to the colony. Ants foraging on tough leaves were larger than ants foraging on soft leaves, and there was a significant reduction in forager size for both plants when the colony was exposed to Neodohrniphora sp. However, there were no relative differences on forager size between the two plants after the introduction of the parasitoid. The lack of response of Neodohrniphora sp. to the increase in ant size when the colony was given tough leaves may be attributed to the unusually large number of suitable hosts in a laboratory colony. However, large foragers are much less abundant in the field, in which case shifts in the size of the workforce triggered by different substrates could affect the incidence of parasitism.     

Interactions with Combined Chemical Cues Inform Harvester Ant Foragers' Decisions to Leave the Nest in Search of Food

Social insect colonies operate without central control or any global assessment of what needs to be done by workers. Colony organization arises from the responses of individuals to local cues. Red harvester ants (Pogonomyrmex barbatus) regulate foraging using interactions between returning and outgoing foragers. The rate at which foragers return with seeds, a measure of food availability, sets the rate at which outgoing foragers leave the nest on foraging trips. We used mimics to test whether outgoing foragers inside the nest respond to the odor of food, oleic acid, the odor of the forager itself, cuticular hydrocarbons, or a combination of both with increased foraging activity. We compared foraging activity, the rate at which foragers passed a line on a trail, before and after the addition of mimics. The combination of both odors, those of food and of foragers, is required to stimulate foraging. The addition of blank mimics, mimics coated with food odor alone, or mimics coated with forager odor alone did not increase foraging activity. We compared the rates at which foragers inside the nest interacted with other ants, blank mimics, and mimics coated with a combination of food and forager odor. Foragers inside the nest interacted more with mimics coated with combined forager/seed odors than with blank mimics, and these interactions had the same effect as those with other foragers. Outgoing foragers inside the nest entrance are stimulated to leave the nest in search of food by interacting with foragers returning with seeds. By using the combined odors of forager cuticular hydrocarbons and of seeds, the colony captures precise information, on the timescale of seconds, about the current availability of food.     

Colony and Individual Forager Responses to Food Quality in the New World Bumble Bee, Bombus occidentalis

The ability of a successful forager to activate colony foraging allows colonies to rapidly exploit ephemeral resources and is an important innovation in the evolution of sociality. We tested the ability of the species, Bombus occidentalis, to stimulate colony foraging for food varying in quality. We then analyzed the behavior of successful foragers inside the nest to learn more about potential foraging activation movements. The number of bees entering a foraging arena was positively correlated with food sucrose concentration (0.5, 1.0, and 2.5 M sucrose, equal to 16–65% w/w). Foragers spent significantly more time imbibing higher concentration solutions. Foragers then returned to the nest where they moved in elaborate paths at variable speeds. There was no significant effect of sucrose concentration on average forager velocity or time spent inside the nest. However, the length of a forager’s path inside the nest (total of all distances moved each 0.1 s) significantly increased with sucrose concentration. On average, individuals foraging on 2.5 M and 1.0 M solution walked paths respectively 1.6 fold and 1.4 longer than the paths of individuals foraging on 0.5 M solution. These longer paths could result in a greater number of nestmate contacts, a factor shown to be important in the activation of B. impatiens foragers and also reported in B. terrestris foragers.     

Independent colony founding by ergatoid queens in the ant genus Pogonomyrmex: queen foraging provides an alternative to dependent colony founding

Ant queens exhibit two primary strategies to initiate nests, independent colony founding (ICF) by solitary queens and dependent colony founding (DCF) when the queen starts a nest with a group of workers that disperse on foot from the parent nest. Numerous ant species have wingless (ergatoid) queens, and it is generally assumed that these species exhibit obligate DCF because their lack of wing musculature provides them with few resources to divert towards producing their first brood of workers. Thus, ICF by ergatoid queens is viewed as maladaptive because these queens need to take additional dangerous foraging trips to garner sufficient food to rear their first brood of workers. Contrary to this prediction, I document ICF by ergatoid queens for three species of harvester ants in the genus Pogonomyrmex (subfamily Myrmicinae), P. cunicularius cunicularius, P. cunicularius pencosensis, and P. huachucanus. Queens of P. huachucanus were obligate foragers, i.e., no minim workers could be produced without external food, and one queen of P. cunicularius pencosensis was observed foraging in the field. Abundant and/or predictable food resources likely select for the evolution of semi-claustral nest founding and ICF by these ergatoid queens. Under these conditions, foraging time would be minimized and the number and size of minim workers would be maximized. These benefits should increase founding success, which could compensate for loss of long-range dispersal. Overall, this study demonstrates that care should be taken before concluding that ant colonies employ DCF based solely on queen morphology.     

Onset of morning activity in bumblebee foragers under natural low light conditions

Foraging on flowers in low light at dusk and dawn comes at an additional cost for insect pollinators with diurnal vision. Nevertheless, some species are known to be frequently active at these times. To explore how early and under which light levels colonies of bumblebees, Bombus terrestris, initiate their foraging activity, we tracked foragers of different body sizes using RFID over 5 consecutive days during warm periods of the flowering season. Bees that left the colony at lower light levels and earlier in the day were larger in size. This result extends the evidence for alloethism in bumblebees and shows that foragers differ in their task specialization depending on body size. By leaving the colony earlier to find and exploit flowers in low light, larger‐sized foragers are aided by their more sensitive eyes and can effectively increase their contributions to the colony''s food influx. The decision to leave the colony early seems to be further facilitated by knowledge about profitable food resources in specific locations. We observed that experience accrued over many foraging flights determined whether a bee started foraging under lower light levels and earlier in the morning. Larger‐sized bees were not more experienced than smaller‐sized bees, confirming earlier observations of wide size ranges among active foragers. Overall, we found that most foragers left at higher light levels when they could see well and fly faster. Nevertheless, a small proportion of foragers left the colony shortly after the onset of dawn when light levels were below 10 lux. Our observations suggest that bumblebee colonies have the potential to balance the benefits of deploying large‐sized or experienced foragers during dawn against the risks and costs of foraging under low light by regulating the onset of their activity at different stages of the colony''s life cycle and in changing environmental conditions.     

Foraging decisions of individual workers vary with colony size in the greenhead ant Rhytidoponera metallica (Formicidae, Ectatomminae)

Summary. The ability of worker ants to adapt their behaviour depending on the social environment of the colony is imperative for colony growth and survival. In this study we use the greenhead ant Rhytidoponera metallica to test for a relationship between colony size and foraging behaviour. We controlled for possible confounding ontogenetic and age effects by splitting large colonies into small and large colony fragments. Large and small colonies differed in worker number but not worker relatedness or worker/brood ratios. Differences in foraging activity were tested in the context of single foraging cycles with and without the opportunity to retrieve food. We found that workers from large colonies foraged for longer distances and spent more time outside the nest than foragers from small colonies. However, foragers from large and small colonies retrieved the first prey item they contacted, irrespective of prey size. Our results show that in R. metallica, foraging decisions made outside the nest by individual workers are related to the size of their colony.Received 23 March 2004; revised 3 June 2004; accepted 4 June 2004.     

Forager size and ecology of Acromyrmex coronatus and other leaf-cutting ants in Costa Rica

I compare forager size and foraging ecology of the leaf-cutting ant Acromyrmex coronatus (Fabricius) with published data on three other leaf-cutter species in Costa Rica, Atta cephalotes (L.), Acromyrmex octospinosus (Reich), and Acromyrmex volcanus Wheeler. Intra-and interspecific differences in forager size in these leaf-cutting ants appear to reflect the economics of harvesting different preferred resources. Ac. coronatus colonies have relatively small foragers (mean mass=3.4±1.4 mg) that cut almost exclusively the thin, soft leaves and other parts of small herbaceous plants. Similarly, small A. cephalotes colonies have small foragers (3.3±1.0 mg) that attack the leaves of small herbaceous plants. In contrast, mature A. cephalotes colonies have a wider sizerange of foragers (7.3±4.1 mg) that primarily attack the leaves of trees, with larger foragers cutting thicker, tougher leaves. In A. cephalotes, the match of forager size to leaf type (both ontogenetically and behaviorally) increases foraging efficiency. Extreme forager polymorphism in mature A. cephalotes colonies appears to broaden the diversity of tree species that they can exploit efficiently. Ac. octospinosus and Ac. volcanus both have large, relatively monomorphic foragers (13.3±4.2 mg and 30.6±4.3 mg, respectively) that typically scavenge for pieces of fallen vegetation, such as dead leaves, fruit, and flowers, in addition to cutting herbs. The large foragers of Ac. octospinosus and Ac. volcanus appear to be well suited as generalist foragers, able to cut or collect any desirable vegetation encountered. Ac. coronatus is similar to A. cephalotes in other ways. Both Ac. coronatus and A. cephalotes establish and maintain cleared trunk trails for foraging, and both have minima workers that hitchhike on the loads carried by foragers, apparently serving to protect the larger foragers from attack by phorid flies. Trunk trails and hitchhikers are not known for Ac. octospinosus and Ac. volcanus. That A. coronatus and A. cephalotes show little overlap in geographic distribution within Costa Rica may relate both to differences in habitat requirements and to interspecific competition.     

Temperature and forager body size affect carbohydrate collection in German yellowjackets, Vespula germanica (Hymenoptera, Vespidae)

The classic formulation of optimal foraging theory predicts that a central-place forager will gather more food if it is required to travel farther from the nest to find that food. We examined the foraging behavior of German yellowjackets (Vespula germanica) to determine whether carbohydrate foragers follow this pattern. We trained foragers to collect 2 M fructose solution at 5 or 50 m from the nest and measured the time spent feeding, load size, and the rate of delivery. We show that as a forager’s crop fills during a foraging bout, the amount of solution ingested per second decreased. However, load size did not change as wasps collected food up to 50 m from the nest. Instead, temperature and body size were better predictors of the volume of fructose a forager carried. Finally, the rate of fructose delivered to the nest was higher at warmer temperatures. Due to the fact that wasps gather more food but feed for shorter periods of time at warmer temperatures, we found an overall negative relationship between feeding time and load size. We conclude that the strong effects temperature had on the behavior of V. germanica foragers imply that feeding time may not always be an accurate predictor of the size of the load an individual carries back to the nest. Results from this study suggest that in yellowjacket colonies, foragers can collect and bring disproportionately more food back to the nest during the warmest days of the summer, a time of year when this pest species reaches peak population size during its annual colony cycle.     

Foraging energetics of the ant,Paraponera clavata

The energy currencies used by foraging animals are expected to relate to the energy costs and benefits of resource collection. However, actual costs of foraging are rarely measured. We measured the ratio of energetic benefit relative to cost (B/C) during foraging for the giant tropical ant, Paraponera clavata. The B/C ratio was 3.9 for nectar-foragers and 67 for insect prey foragers. In contrast, the B/C ratio during foraging for seed harvester ants (Pogonomyrmex occidentalis) is over 1000, demonstrating that the B/C ratio can vary widely among ants. The B/C ratio was 300 times lower for nectar-foraging Paraponera than for the seed-harvesting Pogonomyrmex because of: (1) a 5-fold lower energetic benefit per trip, (2) a 10-fold greater cost due to longer foraging distances, and (3) a 6-fold greater energy cost per meter due to larger body size. For Paraponera daily colonial energy intake rates are similar to expeditures and may limit colony growth and reproduction. In contrast, for Pogonomyrmex energy intake rates are an order of magnitude higher than estimated costs, suggesting that Pogonomyrmex colonies are unlikely to be limited by short-term energy intake. We suggest that variation in individual B/C ratios may explain why the foraging behavior of Paraponera but not Pogonomyrmex appears sensitive to foraging costs.     

Size variation and foraging rate in bumblebees (Bombus terrestris)

Summary: Size polymorphism is an important life history trait in bumblebees with strong impact on individual behavior and colony organization. Within a colony larger workers tend to serve as foragers, while smaller workers fulfill in-hive tasks. It is often assumed that size-dependent division of labor relates to differences in task performance. In this study we examined size-dependent interindividual variability in foraging, i.e. whether foraging behavior and foraging capability of bumblebee workers are affected by their size. We observed two freely foraging Bombus terrestris colonies and measured i) trip number, ii) trip time, iii) proportion of nectar trips, and iv) nectar foraging rate of different sized foragers. In all observation periods large foragers exhibited a significantly higher foraging rate than small foragers. None of the other three foraging parameters was affected by worker size. Thus, large foragers contributed disproportionately more to the current nectar influx of their colony. We provide a detailed discussion of the possible proximate mechanisms underlying the differences in foraging rate.     

Colony variation in the collective regulation of foraging by harvester ants

This study investigates variation in collective behavior in a natural population of colonies of the harvester ant, Pogonomyrmex barbatus. Harvester ant colonies regulate foraging activity to adjust to current food availability; the rate at which inactive foragers leave the nest on the next trip depends on the rate at which successful foragers return with food. This study investigates differences among colonies in foraging activity and how these differences are associated with variation among colonies in the regulation of foraging. Colonies differ in the baseline rate at which patrollers leave the nest, without stimulation from returning ants. This baseline rate predicts a colony's foraging activity, suggesting there is a colony-specific activity level that influences how quickly any ant leaves the nest. When a colony's foraging activity is high, the colony is more likely to regulate foraging. Moreover, colonies differ in the propensity to adjust the rate of outgoing foragers to the rate of forager return. Naturally occurring variation in the regulation of foraging may lead to variation in colony survival and reproductive success.     

Division of labour and specification of castes in the red imported fire ant Solenopsis invicta buren

Division of labour in Solenopsis invicta follows a familiar pattern: younger, smaller ants tend toward brood care while older, larger ants tend toward foraging. However, long-term observations of marked individuals reveal that length of nursing and foraging ‘careers’ and the age of transition between these activities vary considerably between and within size groups, and are related to length of life. Experiments with entire colonies show that larger ants are more likely than smaller ants to forage for insect prey. There are two main worker castes, ‘nurses’ and ‘foragers’, whose members span a wide age-size range, and a large ‘reserve’ subcaste, heterogeneous in age, size, and behaviour: reserves may nurse, forage, store liquid food, or relay food from nurses to foragers. The proportion of ants engaged in foraging decreases with colony size because many ants in large colonies are not exposed to recruitment signals.     

Diet of two sympatric Pheidole spp. ants in the central Monte desert: implications for seed–granivore interactions

Ants of the genus Pheidole are important seed consumers in several desert ecosystems. In South American deserts, although several Pheidole spp. have been characterized as seed harvesters, studies on their diet and ecological role are still missing. Pheidole spininodis (Mayr) and Pheidole bergi (Mayr) are capable of removing seeds in the central Monte desert. The aim of this study was to quantify and compare the diet of these species and to interpret the results in the context of seed–granivore interactions. Diet was estimated during mid-summer by collecting items brought back to the nest by foragers in ten colonies per species. While P. spininodis was mainly granivorous, P. bergi was mainly insectivorous. However, they both collected ~40% of other types of items. Among seeds, the diet of P. spininodis included mostly grass seeds, whereas the diet of P. bergi was mainly made up of shrub and tree seeds, usually retrieved cooperatively. This behavior allowed P. bergi to carry larger seeds, resulting in diet partitioning in terms of seed size. However, diet of P. spininodis is very similar to that of three sympatric Pogonomyrmex species. Thus, specialized harvester ants remove large quantities of grass seeds in the central Monte desert during the summer, potentially affecting their abundance in the soil seed bank. P. bergi directs its feeding pressure to shrub and tree seeds, and although seeds constitute ~10% of its diet, its high colony density and high activity levels, added to the lower proportion of large seeds in the soil seed bank, indicate that their importance as seed consumers cannot be ruled out.     

Polymorphism and size-pairing in the harvester antPogonomyrmex badius: a test of the ecological release hypothesis

Summary The role of caste polymorphism in the foraging strategy ofPogonomyrmex badius was studied in the field by measuring food items collected by foragers, and correlating food item size variables with forager size variables. The diet ofP. badius included seeds and insects. In two colonies examined, these food types comprised different proportions of the diet sample.Although some forager size variables showed close or significant correlations with food item size variables, we could identify no overall significant relationship between worker size and seed or prey size. Polymorphism inP. badius may be associated with omnivory. However, since minor workers serve as foragers and represent a portion of the total worker size variation, dietary expansion through caste proliferation appears to be only one aspect of the functional significance of polymorphism in this species.