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.     

Ant diversity and its relationship with vegetation and soil factors in an alluvial fan of the Tehuacán Valley, Mexico

In this study, we analyze the ant community found along an alluvial fan located in the Tehuacán Valley, central Mexico. Considering that this fan is composed of four terraces with different soils and vegetation structures, our main goal was to determine whether there are significant differences in ant diversity among terraces. To accomplish this goal, we determine species richness and abundance in order to calculate diversity and evenness indices. In addition, we classify species in different feeding guilds to evaluate whether differences among terraces exist. We expected higher ant diversity and variety of food guilds in terraces with sandy soils and complex vegetation structures than in terraces with argillic and calcic horizons. Correlations between several diversity parameters, and soil percent-sand and vegetation structure were also conducted. A total of 26 ant species were recorded along the fan. Species richness was not different among terraces whereas abundance was higher in sandy soils and on terraces with complex vegetation structure. Particularly, the abundance of the harvester ant Pogonomyrmex barbatus was higher in these terraces decreasing total ant diversity and evenness. Species richness within feeding guilds was similar among terraces with the generalized foragers as the most common. Our work suggests that percentage of sand in the soil and complexity of vegetation structure of the alluvial fan studied might be influencing ant distribution and favoring the abundance of numerically dominant species which could be affecting the diversity patterns of the whole community.     

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.     

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.     

Seasonal effects of granivory on broadcast seeding restoration in a degraded semiarid grassland

Granivores, such as rodents and harvester ants, are common in dryland ecosystems. In intact dryland ecosystems, the effects of such granivores on soil seed banks, plant community structure, and ecosystem functioning are well established. However, the effects of granivores on dryland restoration have received much less attention. In this study, we evaluated the seasonality of seed removal by harvester ants, seed preference of harvester ants, and effects of granivore exclosure on the soil seed bank at four sites in southeastern Utah. We studied two species of harvester ants, Pogonomyrmex rugosus and P. occidentalis. The foraging behavior of Pogonomyrmex spp. was temperature-dependent; they removed no seeds during winter months, and up to 80% of seeds during high activity in the summer. Despite seasonal differences in seed removal, Pogonomyrmex spp. did not show significant preferences for seed species. However, their seasonal foraging behavior translated into measurable differences in an experimental soil seed bank. Over the summer months, ants reduced the number of seeds in the seed bank by approximately 20%, but when ants were excluded during the winter, the number of seeds in the seed bank remained the same. This finding supports the timing of business-as-usual seeding practices of applying seeds in late fall. If seeds germinate in the first season after application, ant granivory is unlikely to pose a large barrier to establishment. However, if seeds remain dormant during the first year, as is common in many dryland species, seeds are vulnerable to ant granivory.     

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.     

Alarm-recruitment behaviour in Pheidole militicida (Hymenoptera: Formicidae)

ABSTRACT.

• 1 Pheidole militicida Wheeler, a seed-harvesting species of the southwestern United States, possesses a major worker caste (soldiers) with unusually large heads. Previous work suggested that these large major workers are specialized defenders against large seed-harvesting species in the genus Pogonomyrmex.
• 2 Experimental introductions of Pogonomyrmex maricopa Wheeler and the omnivorous fire ant, Solenopsis xyloni McCook, demonstrate that P. militicida colonies alarm-recruit major workers against S.xyloni workers but not against the larger Pogonomyrmex workers.
• 3 P.militicida is the second species in which enemy-specific defence against Solenopsis fire ants has been detected. I suggest that the special role of Solenopsis as both competitor and predator in ant communities frequently favours major worker specialization against this genus.
• 4 Major workers of P.militicida and Pheidole dentata Mayr, the other species with enemy-specific defence against Solenopsis, are morphologically very different from one another. It is suggested that interspecific variation in major worker morphology is not necessarily associated with variation in behavioural specialization.

     

Individual foraging components of harvester ants: movement patterns and seed patch fidelity

Summary We investigated individual foraging components of the western harvester ant,Pogonomyrmex occidentalis, in the native seed background of a shrub-steppe environment. Our study identified factors affecting foraging movements and seed selection by individual ants. Some assumptions and predictions of central-place foraging theory and a correlated random walk were evaluated for individual foragers. Results showed that ant size was only weakly correlated with the seed sizes harvested; seed size was a more important constraint than a predictor of seed selection. Individual ants spent more time in localized search behavior than traveling between search areas and nests.P. occidentalis foragers encountered seeds randomly with respect to time, and handled a mean of 1.7 seeds/trip. A correlation of increased search effort with greater travel distances was consistent with central-place foraging theory but, contrary to it, search and travel effort were not associated with energetic reward.Individual ants exhibited fidelity in both search site and native seed species. Spatial analyses of foraging movements showed a highly oriented travel path while running, and an area-restricted path while searching. Searching ants moved in a manner consistent with a correlated random walk. The deterministic component of patch fidelity and the stochastic component of search may override energetic foraging decisions in individualP. occidentalis ants.     

SEXUAL SELECTION ON BODY SIZE AND SHAPE IN THE WESTERN HARVESTER ANT,POGONOMYRMEX OCCIDENTALIS CRESSON

Mating in social insects has generally been studied in relation to reproductive allocation and relatedness. Despite the tremendous morphological diversity in social insects, little is known about how individual morphology affects mating success. We examined the correlation of male size and shape with mating success in the western harvester ant, Pogonomyrmex occidentalis. Larger males had significantly higher mating success in two independent collections of males at mating aggregations. We also detected significant linear and nonlinear selection on aspects of male shape that were consistent across years. These shape components are independent of size, suggesting that male mating success is a complex function of size and shape. Successful males had elongate thoraxes and short mandibles relative to males collected at random at the lek. Overall, mated males also had longer postpetioles relative to body size, but there was also evidence of nonlinear selection on relative postpetiole length in both years. We found no evidence of assortative mating based on size or multivariate shape measures in either year, but in one year we found weak assortative mating based on some univariate traits.