Directional fidelity as a foraging constraint in the western harvester ant,Pogonomyrmex occidentalis

Summary Understanding the foraging behavior of an animal is critically dependent upon knowledge of the constraints on that animal. In this study, I tested whether fidelity to foraging direction acts as a behavioral constraint to foraging western harvester ants, Pogonomyrmex occidentalis. Individual P. occidentalis foragers showed strong fidelity to foraging route and direction. Directional fidelity in this population was not related to trunk trail use, food specialization, colony activity levels, or mortality risks. Directional fidelity constrained individual foraging decisions; when colonies were offered seeds of different quality in 2 directions, individuals did not switch directions to obtain the energetically more rewarding seeds. Colony-level recruitment was increased for energetically more profitable seeds, indicating that colonial responses may compensate for the constraints of directional fidelity on individual foragers.     

Recruitment pheromone in the harvester ant genus Pogonomyrmex

Workers of the harvester ant genus Pogonomyrmex employ recruitment trail pheromones discharged from the poison gland. In P. barbatus, P. maricopa, P. occidentalis and P. rugosus we identified three pyrazines [2,5-dimethylpyrazine, trimethylpyrazine and 3-ethyl-2,5-dimethylpyrazine (EDMP)] as major compounds of the volatile part of the poison-gland secretions. Laboratory and field tests revealed EDMP to be the main recruitment pheromone.     

Characterization of polymorphic microsatellite loci in the red harvester ant,Pogonomyrmex barbatus

The red harvester ant, Pogonomyrmex barbatus, is a monogynous, polyandrous species: each ant colony is founded by a single queen that has mated with one or more males. To study levels of polyandry within a colony, as well as relationships among colonies, we developed 10 polymorphic microsatellite loci for P. barbatus. With the number of alleles per locus ranging from 3 to 39, and expected heterozygosities of 0.58–0.95, these markers promise to be useful in the study of colony and population genetic structure.     

Male size, sperm transfer, and colony fitness in the western harvester ant, Pogonomyrmex occidentalis

Mating success in the western harvester ant, Pogonomyrmex occidentalis, increases with male size. We tested the hypothesis that increased mating success increases male fitness and the fitness of colonies that make large males by comparing the sperm content of males prior to and at the conclusion of the mating swarm. The number of sperm a male initially possesses is a function of male size, and large males transfer a greater proportion of their sperm than do small males. For colonies, the payoff per unit of investment is an increasing function of male size, and investment in large males is not equivalent to investing in a larger number of small males. Allocation ratios in species that show size variation in reproductives may need to be modified by the individual fitness functions.     

The adaptive nature of non-food collection for the Florida harvester ant, Pogonomyrmex badius

Abstract.  1. Many animals forage for non-food objects, such as nesting materials. In many organisms, the depots of such objects can be expansive, and are not intuitively adaptive. This study empirically explores varying theories for why these depots exist in the Florida harvester ant, Pogonomyrmex badius .
2. The Florida harvester ant, like its western congeners, collects and deposits large quantities of non-food objects around its nest entrance. The vast majority of objects sampled from colonies in this study population were charcoal pieces whereas most western species collect pebbles.
3. Objects in these depots can number in the millions, suggesting a large cost to this behaviour and strengthening an adaptationist argument.
4. Many hypotheses have been suggested for these depots, including temperature and water modification and territoriality. No evidence was found to support the results of a previous study which found that charcoal depots deter other ant species.
5. The depots do not appear to affect soil hydrology, but do slightly increase soil temperature at the very top of the nest. In an attempt to estimate the seasonal fitness of colonies with and without charcoal depots, no statistical difference was detected in sexual investment between treatment groups.
6. These results suggest that the charcoal depots may be selectively advantageous through temperature modification, but may serve alternate functions that are species and habitat specific.     

Diet and seed removal rates by the harvester ants Pogonomyrmex rastratus and Pogonomyrmex pronotalis in the central Monte desert, Argentina

Granivorous animals, through seed consumption, may have an important influence on plant abundance, distribution and species composition in desert ecosystems. The aims of this study are twofold: to quantify the diet of Pogonomyrmex rastratus (Mayr) and pogonomyrmex pronotalis (Santschi), and to estimate seed removal per colony of both species throughout their activity season (October-April) in the central Monte desert, Argentina. Both species rely heavily upon seeds, which account for 87–94 % of the items carried to the nests. Their diets are similar, consisting mainly of grass seeds, which represent more than 93 % of the seeds. Among them, three species predominate: Aristida spp., Trichloris crinita and Pappophorum spp. Seasonal variations as well as seed species richness in the diet are also similar between species. However, their food-handling behaviour differs: most caryopses carried by P. pronotalis bear bracts whereas most caryopses carried by P. rastratus lack them. Seed removal per colony by P. rastratus (6 × 10⁴ seeds/colony) and by P. pronotalis (5 × 10⁴ seeds/colony) throughout the season is similar to the one reported for P. occidentalis in North America. However, seed removal per hectare, which could be estimated for P. rastratus (8.3 × 10⁵ seeds/ha), is lower than removal rates reported for the North American species P. barbatus, P. desertorum, P. rugosus and P. californicus, probably because P. rastratus has lower activity levels and smaller colonies than the North American studied species. Received 6 April 2005; revised 1 September 2005; accepted 2 September 2005.     

Characterization and distribution of Pogonomyrmex harvester ant lineages with genetic caste determination

Genetic caste determination has been described in two populations of Pogonomyrmex harvester ants, each comprising a pair of interbreeding lineages. Queens mate with males of their own and of the alternate lineage and produce two types of diploid offspring, those fertilized by males of the queens' lineage which develop into queens and those fertilized by males of the other lineage which develop into workers. Each of the lineages has been shown to be itself of hybrid origin between the species Pogonomyrmex barbatus and Pogonomyrmex rugosus, which both have typical, environmentally determined caste differentiation. In a large scale genetic survey across 35 sites in Arizona, New Mexico and Texas, we found that genetic caste determination associated with pairs of interbreeding lineages occurred frequently (in 26 out of the 35 sites). Overall, we identified eight lineages with genetic caste determination that always co-occurred in the same complementary lineage pairs. Three of the four lineage pairs appear to have a common origin while their relationship with the fourth remains unclear. The level of genetic differentiation among these eight lineages was significantly higher than the differentiation between P. rugosus and P. barbatus, which questions the appropriate taxonomic status of these genetic lineages. In addition to being genetically isolated from one another, all lineages with genetic caste determination were genetically distinct from P. rugosus and P. barbatus, even when colonies of interbreeding lineages co-occurred with colonies of either putative parent at the same site. Such nearly complete reproductive isolation between the lineages and the species with environmental caste determination might prevent the genetic caste determination system to be swept away by gene flow.     

Effect of harvester ants of the genus Pogonomyrmex on the soil seed bank around their nests in the central Monte desert,Argentina

1. The abundance and composition of soil seed banks is a key determinant of plant community structure. Harvester ants can remove huge quantities of preferred seeds close to the nest affecting composition and spatial distribution of plants. 2. In the central Monte desert (Argentina) ants of the genus Pogonomyrmex have high seed removal rates, especially of the five main grasses. The aim of this study was to establish if their foraging activity affects spatial patterns of the soil seed bank around their nests. Our hypotheses were: (1) removal by ants decreases seed abundance of preferred species in the soil; and (2) the effect varies in time. 3. Soil seed abundance was assessed at different distances from Pogonomyrmex nests in the litter and in bare soil at the beginning, the middle, and the end of the season (late spring‐early autumn). 4. A lower seed abundance of preferred species was observed close to the nest in the litter at the end of the season. Non‐preferred species showed no distance gradient. 5. The lower foraging activity and seed consumption at the beginning of the season could explain the temporal variation of the spatial effect. This was only observed in the litter, probably because of the higher removal frequency in this substrate. 6. Colonies of Pogonomyrmex spp. could enhance the heterogeneity of soil seed banks in the central Monte desert from the summer to the beginning of the autumn. Implications for vegetation dynamics depend on the degree to which seed density limits perennial grasses recruitment after ant activity season.     

Genetic caste determination in Pogonomyrmex harvester ants imposes costs during colony founding

Some populations of Pogonomyrmex harvester ants comprise genetically differentiated pairs of interbreeding lineages. Queens mate with males of their own and of the alternate lineage and produce pure-lineage offspring which develop into queens and inter-lineage offspring which develop into workers. Here we tested whether such genetic caste determination is associated with costs in terms of the ability to optimally allocate resources to the production of queens and workers. During the stage of colony founding, when only workers are produced, queens laid a high proportion of pure-lineage eggs but the large majority of these eggs failed to develop. As a consequence, the number of offspring produced by incipient colonies decreased linearly with the proportion of pure-lineage eggs laid by queens. Moreover, queens of the lineage most commonly represented in a given mating flight produced more pure-lineage eggs, in line with the view that they mate randomly with the two types of males and indiscriminately use their sperm. Altogether these results predict frequency-dependent selection on pairs of lineages because queens of the more common lineage will produce more pure-lineage eggs and their colonies be less successful during the stage of colony founding, which may be an important force maintaining the coexistence of pairs of lineages within populations.     

Molecular evidence for the origin of workerless social parasites in the ant genus Pogonomyrmex

Abstract.— Speciation of two social parasites from their respective hosts is tested using a molecular phylogeny. Alignment of 711 DNA base pairs of mitochondrial cytochrome b gene was used to assess phylogenetic relationships of inquiline species to their hosts and to other members of the genus. We show that the inquiline social parasites of the North American seed harvester ants are monophyletic, descending from one of the known hosts ( Pogonomyrmex barbatus ) in the recent past and shifting hosts in a pattern similar to that observed in other Hymenopteran social parasites. In addition, the host populations unexpectedly were found to be polyphyletic. Populations of Pogonomyrmex rugosus from an area east of the Chiricahua Mountains in Southern Arizona belong to a mitochondrial clade separate from the more western clade of P. rugosus from the Sonoran and Chihuahuan Deserts. Evidence of mitochondrial DNA introgression between P. rugosus and P. barbatus was also observed. We conclude that Emery's rule does not strictly hold for this system, but that the hosts and parasites are very closely related, supporting a loose definition of Emery's rule.     

The Old Ladies of the Seed Harvester ant Pogonomyrmex Rugosus: Foraging Performed by Two Groups of Workers

We examined temporal polyethism in Pogonomyrmex rugosus, predicting a pattern of decreasing age from foragers to nest maintenance workers to individuals that were recruited to harvest a temporary food source. Nest maintenance workers were younger than foragers, as indicated by their heavier mass and lower mandibular wear. In contrast, recruited foragers were similar in mass to foragers but they displayed higher mandibular wear, suggesting that they were at least as old as foragers. Longevity estimates for marked individuals of these two latter task groups showed mixed results. Higher mandibular wear of recruited foragers suggests that they did not follow the normal sequence for temporal polyethism, but rather that they functioned as seed-millers, which should more quickly abrade their dentition. This would be the first demonstration of specialist milling individuals in a monomorphic seed-harvester ant.     

Influence of seed size and seed nature on recruitment in the polymorphic harvester ant Messor barbarus

In the polymorphic harvester ant Messor barbarus, the colony modulates response during foraging according to seed characteristics. The easiness of picking up small seeds (oat fragments or canaryseeds) shortens the time lapse between food discovery and the return of scouts to the nest, favouring the onset of recruitment and higher mobilisation rates than bigger seeds (whole oat). However, size is not the only criterion, as canaryseeds are preferred among small ones. A modulation of the trail laid by the first scouts also plays a role in the shaping of harvesting patterns and in the enhancement of recruitment towards small and/or preferred seed. This flexibility extends to the relative participation of the three worker sizes classes, which differs according to seed. Media are the most numerous at the foraging arena and largely responsible for trail-laying. Minor participate in seed harvesting and trail-laying, mostly when they can be efficient in carrying seeds, such as oat fragments. Major scarcely participate in harvesting and trail-laying, but are involved in the exploitation of bigger and/or preferred seed species. When faced with different seed species, M. barbarus seems thus to collectively adopt a “foraging time minimisation” rather than an “energetic gain per item harvested maximisation” strategy. Dynamics of foraging and division of labour between foragers of different sizes are related to both the workers’ ability to carry seed items, and the transfer of information to nestmates by trail-laying, all parameters that vary according to seed characteristics.     

Polyandry and fitness in the western harvester ant, Pogonomyrmex occidentalis

Using four highly polymorphic microsatellite markers (12-28 alleles), we gentoyped workers from 63 colonies of Pogonomyrmex occidentalis. Colonies have a single, multiply mated queen, and an average number of 6.3 patrilines per colony. Colony growth was measured over an 8-year period in the study population. Intracolonial relatedness and colony growth are correlated negatively, indicating a substantial fitness benefit to multiple mating.     

Density-dependent foraging in the harvester ant Messor ebeninus: two experiments

Harvester ants are important seed predators in many xeric environments, and their foraging choices can influence the composition of plant communities. Seed abundance has been cited as an important factor in determining such foraging preferences. Three seed types (sesame, millet, and flax) were experimentally introduced in differing proportions near nests of the ant Messor ebeninus near the Dead Sea, in territory administered by the state of Israel. Two experiments were designed to investigate the effects of this density conditioning on two stages of the ants' subsequent foraging behavior: recruitment to seed patches and selection of seeds from within a patch. When seeds were presented in small, single-species patches, experimentally common seeds were discovered and exploited significantly faster than rare seeds, especially among less preferred seed types. When seeds were presented in large, mixed patches, however, no consistent effect of density was observed.     

Influence of temporal fluctuations in seed abundance on the foraging behaviour of harvester ants (Pogonomyrmex spp.) in the central Monte desert,Argentina

The simultaneous study of the temporal dynamics of foraging behaviour, diet and seed abundance is essential to assess the way in which resources affect the behaviour and ecology of harvester ants. Here, we evaluate how fluctuations in grass seed abundance during three consecutive growing seasons influenced the foraging behaviour and diet of the harvester ants Pogonomyrmex rastratus, P. mendozanus and P. inermis in the central Monte desert, Argentina. Seed abundance of the most consumed grasses varied greatly through ant activity season, and ants altered their foraging behaviour in response to those changes. Foragers spent more time travelling and searching for food, and their foraging trips took longer during the low seed availability season. Foraging distance was very similar among species and, contrary to our expectations, did not vary between seasons. Foraging success of P. rastratus and P. inermis increased during the high availability season. This matched the seasonal pattern of foraging activity, suggesting that colonies may detect seed abundance and regulate their foraging effort with the rate of forager success. Although grass seeds were the main item in the diet of the three species, P. mendozanus, and to a lesser extent P. rastratus, turned more generalist when grass seeds were scarce. In contrast, P. inermis showed a very narrow diet breadth, only harvesting grass seeds in both seasons. Our results indicate the relevance of seed availability on foraging behaviour of harvester ants, which should be taken into account when predicting and evaluating the effect of ants on seed resources as well as numerical responses of harvester ant populations to the temporal and spatial variations in grass seed abundance.     

Two alternate mechanisms contribute to the persistence of interdependent lineages in Pogonomyrmex harvester ants

Some populations of Pogonomyrmex harvester ants comprise pairs of highly differentiated lineages with queens mating at random with several males of their own and of the alternate lineage. These queens produce two types of diploid offspring, those fertilized by males of the queens' lineage which develop into new queens and those fertilized by males of the other lineage which mostly develop into functionally sterile workers. This unusual mode of genetic caste determination has been found in 26 populations and a total of four lineage pairs (F(1)-F(2), G(1)-G(2), H(1)-H(2) and J(1)-J(2)) have been described in these populations. Despite the fact that a few interlineage queens are produced, previous studies revealed that there is a complete lack of genetic introgression between lineages. Here we quantify the proportion of interlineage queens produced in each of the four lineage pairs and determine the fate of these queens. In the F(1)-F(2), G(1)-G(2) and H(1)-H(2) lineage pairs, interlineage queens were produced by a minority of colonies. These colonies exclusively produced interlineage queens and workers, suggesting that interlineage eggs can develop into queens in these three pairs of lineages in the absence of competition with pure-lineage brood. An analysis of three key stages of the colony life cycle revealed that colonies headed by interlineage queens failed to grow sufficiently to produce reproductive individuals. In laboratory comparisons, interlineage queens produced fewer viable eggs, with the effect that they raised fewer workers and lost more weight per worker produced than pure-lineage queens. In the J(1)-J(2) lineage pair, we did not find a single interlineage queen, raising the possibility that interlineage eggs have completely lost the ability to develop into queens in this lineage pair. Hence, two distinct mechanisms seem to account for the complete lack of between-lineage gene flow in the F(1)-F(2), G(1)-G(2), H(1)-H(2) and J(1)-J(2) lineage pairs.     

Optimal and central-place foraging theory applied to a desert harvester ant,Pogonomyrmex californicus

Summary Certain predictions of optimal- and central place-foraging theory were tested on the desert harvester ant, Pogonomyrmex californicus. Colonies were offered three different sizes of oat seed and found to maximize net energy intake (ei) over time (t _i ) by harvesting the seed sizes with the highest e _i /t _i rank. Two aspects of t _i were measured that were assumed constant in previous studies. The handling components of t _i (time required to manipulate the seed and travel time back to the colony with the food) were measured and found to be positively correlated with seed size. The manipulation success rate (the percentage of handled seeds successfully picked up) decreased with increased seed size. These results point out how important it is to measure all parameters of e _i /t _i rather than to assume constancy with both prey type and foraging distance. The relative abundance of less preferred food types was important in determining the proportion of preferred types in the diet. The food supply of eight colonies was manipulated experimentally over a 25-day period. Four deprived colonies were constrained within aluminum enclosures to prevented foraging. The remaining four satiated colonies were given food ad libitum. The niche breadths of the treated colonies were then compared to controls, but found not to differ significantly. Seed baits were offered at three distances from the colony to test whether selectivity increased with disance. Contrary to theoretical predictions, all colonies harcested about the same proportion of each seed size at each distance.     

Recruitment in the harvester ant, Pogonomyrmex occidentalis: effects of experimental removal

We examined the importance of experimental removal of mature colonies on colony recruitment in the western harvester ant Pogonomyrmex occidentalis. To test the common assumption that established colonies suppress the establishment of new colonies we removed all colonies in ten 0.25 ha plots in 1996 and an additional five plots in 1997 and measured new colony recruitment in 1997, 1998, and 1999. Using a blocked, paired plot design we found that removal of colonies increased new colony recruitment in some areas of the site, but not others. Spatial variation in the importance of established colonies to recruitment was consistent across years; blocks in which density dependence was important in one year exhibited density dependent recruitment in following years. We estimated that in the blocks where recruitment was affected by established colonies, they accounted for less than 10% of the mortality of foundress queens. The increase in the number of new recruits (on average two additional new colonies) was considerably less than the number of colonies removed; average colony density in the removal plots was 14 colonies per 0.25 ha plot. The consistent lack of importance of established colonies to recruitment in one block and the relatively small response to colony removal in the other blocks suggests that the number of new colonies in a year may not be equivalent to the number of deaths of established colonies in that year. Space limitation is an important influence on recruitment in P. occidentalis, but the magnitude of the limitation varies spatially.     

Respiratory physiology and water relations of three species of Pogonomyrmex harvester ants (Hymenoptera: Formicidae)

The respiratory physiology and water relations of three harvester ant species (Pogonomyrmex rugosus Emery, P. occidentalis[Cresson] and P. californicus[Buckley]) were examined at three temperatures (15, 25 and 35°C) using a flow-through respirometry system. As intact ants tended to be active during testing, we performed a parallel set of experiments on individuals rendered motionless by decapitation. Both intact and decapitated ants exhibited discontinuous ventilation. Decapitation caused metabolic rate (V˙CO₂) and burst frequency to decrease in all three species. Burst volume either remained constant or increased after removal of the head, though mass-specific V˙CO₂ was unaffected except in P. rugosus. Mass-specific V˙CO₂s of headless harvesters were comparable with published values derived from motionless specimens of other ant species. The mean Q₁₀ for intact ants of all three species was 2.37; for decapitated insects the mean was 2.32. Respiratory water constituted a small (< 5%) fraction of total loss, and we believe that discontinuous ventilation does not act to conserve water in these organisms, although it may serve other functions.