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.     

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.     

Behavioural flexibility does not prevent numerical declines of harvester ants under intense livestock grazing

1. Human‐induced rapid environmental change may decrease food resources and create unfavourable conditions for native species. Organisms showing a flexible foraging behaviour can exploit novel or alternative foods and are more likely to persist, whereas less flexible organisms might suffer starvation and numerical reductions. 2. This study assessed whether declines in the quality and availability of seeds prompted by grazing provoke behavioural and numerical responses in Pogonomyrmex mendozanus ants, and aimed to test whether behavioural flexibility buffers habitat degradation and prevents numerical declines. 3. Heavy grazing caused seed reductions, especially of the highly consumed and preferred grass seeds, and triggered two kinds of ant responses. Ants expanded their diet by incorporating a greater proportion of non‐seed items, and they reduced foraging activity. As a consequence, the rate of food intake per colony lessened, particularly that of carbohydrate‐rich seeds like grass seeds. Colony density under heavy grazing also fell. 4. Habitat degradation triggered a cascade of mechanisms that starts with a decrease in the seed resources and continues with changes in ant behaviour. However, behavioural responses were insufficient to prevent ant numerical declines. The results of this study suggest that the reduction in the colony density under habitat degradation was provoked by ant nutritional deficit and starvation, and predict a deterioration in body condition, colony performance and reproduction of ants, which deserve further assessment.     

Queen size mediates queen survival and colony fitness in harvester ants

Abstract We examined the effect of queen size on the probability of new colony establishment in the ant Pogonomyrmex occidentalis. Large queens are significantly more likely to survive than small queens through the initial stages of colony founding. These differences in individual fitness correlates have corresponding effects on colony fitness. In species in which individual queens vary in fitness, sexual allocation ratios should incorporate the individual fitness functions.     

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.     

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.     

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.     

Polyandry and Reproduction in the Serpentine Leaf Miner Liriomyza trifolii (Diptera: Agromyzidae)

We investigated the hypotheses that Liriomyza trifolii is a polyandrous species and that multiple mating is required to maximize female fecundity. In addition, the hypothesis that male copulating frequency reduces female fecundity was examined. Using sterile and fertile males, we found that L. trifolii females mate more than once during their life span, and that multiple mating is required to maximize female fecundity. A similar amount of sperm was transferred during five successive male copulations with an equal insemination probability. However, the probability of reproduction failure was strongly correlated with the male copulating sequence. Moreover, the failure rate of sperm transfer was found to be 6.8%. In light of these results, possible selection pressures driving for polyandry in L. trifolii are discussed.     

Soil temperature, digging behaviour, and the adaptive value of nest depth in South American species of Acromyrmex leaf-cutting ants

In leaf-cutting ants, workers are expected to excavate the nest at a soil depth that provides suitable temperatures, since the symbiotic fungus cultivated inside nest chambers is highly dependent on temperature for proper growth. We hypothesize that the different nesting habits observed in Acromyrmex leaf-cutting ants in the South American continent, i.e. superficial and subterranean nests, depend on the occurrence, across the soil profile, of the temperature range preferred by workers for digging. To test this hypothesis, we first explored whether the nesting habits in the genus Acromyrmex are correlated with the prevailing soil temperature regimes at the reported nest locations. Second, we experimentally investigated whether Acromyrmex workers engaged in digging use soil temperature as a cue to decide where to excavate the nest. A bibliographic survey of nesting habits of 21 South American Acromyrmex species indicated that nesting habits are correlated with the soil temperature regimes: the warmer the soil at the nesting site, the higher the number of species inhabiting subterranean nests, as compared to superficial nests. For those species showing nesting plasticity, subterranean nests occurred in hot soils, and superficial nests in cold ones. Experimental results indicated that Acromyrmex lundi workers use soil temperature as an orientation cue to decide where to start digging, and respond to rising and falling soil temperatures by moving to alternative digging places, or by stopping digging, respectively. The soil temperature range preferred for digging, between 20°C and maximally 30.6°C, matched the range at which colony growth would be maximized. It is suggested that temperature-sensitive digging guides digging workers towards their preferred range of soil temperature. Workers’ thermopreferences lead to a concentration of digging activity at the soil layers where the preferred range occurs, and therefore, to the construction of superficial nests in cold soils, and subterranean ones in hot soils. The adaptive value of the temperature-related nesting habits, and the temperature-sensitive digging, is further discussed.     

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.     

Genetic variance of mating frequency in the honeybee (Apis mellifera L.)

Summary. The genetic variance of queen mating frequency was studied in honeybees (Apis mellifera carnica). Worker offspring (N = 966) of 28 naturally mated half sister-queens (r = 0.25) from seven unrelated breeding lines were genotyped at four DNA microsatellites. The mating frequencies of the queens were derived from the offspring genotypes. The number of observed matings per queen ranged from 10 to 28 with an average of 17.32 ± 1.10 (number of estimated matings: 24.94 ± 2.51; number of effective matings: 20.09 ± 1.73). Half-sib analyses of the breeding lines were used to estimate heritability. Heritability was h² = 0.449 ± 0.135 for the estimated number of matings and h² = 0.262 ± 0.103 for the number of effective males, which are both significantly different from zero. We conclude that a high genetic variance for polyandry in honeybees can be favored by balanced selection between individual queen and colony level.Received 16 October 2003; revised 4 May 2004; accepted 4 May 2004.     

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.     

Effects of ants on the foraging of birds in spruce trees

I experimentally excluded ants from randomly selected spruce trees Picea abies near colonies of the wood ant Formica aquilonia. Foraging activity of birds in these trees was then compared to the foraging activity of birds in neighboring spruce trees, where ants were allowed to continue foraging. Birds which foraged in the foliage showed the effects of competition with ants: they visited the trees without ants more frequently, and for longer periods. In addition, the insects and spiders that they utilized as food were more abundant in the foliage of trees without ants. Cone-foraging birds, however, which fed on seeds in cones at the tops of the trees, did not show a preference for trees without ants. The differences of tree usage between foliage-gleaning and coneforaging birds can be explained by alteration of the birds' food supply by wood ants: ants did not feed on seeds in cones, and so did not compete with cone-foraging birds. However, foraging wood ants did feed on arthropods living in the foliage, thus reducing the amount of food available to birds there.     

Population- and sociogenetic structure of the leaf-cutter ant <Emphasis Type="Italic">Atta colombica</Emphasis> (Formicidae,Myrmicinae)

Relatedness and genetic variability in colonies of social insects are strongly influenced by the number of queens present and the number of matings per queen, but also by the genetic variability in the population. Thus, multiple paternity will enhance within-colony genetic variability more strongly when the males a queen mates with are unrelated. To study the kin-structure within colonies of the leaf-cutter ant Atta colombica and the population structure of this species around Barro Colorado Island, Panama, we developed five polymorphic microsatellite loci with a range of three to 17 alleles in At. colombica, all of which cross-amplify in other higher attines as well. The average effective mating frequency calculated from four-locus microsatellite genotypes was 1.89 ± 0.12 (harmonic mean ± SE) and thus slightly lower than the average observed mating frequency of 2.50 ± 0.11 (arithmetic mean ± SE) over the 55 colonies studied, confirming former studies that utilized fewer loci. The discrepancy between observed mating frequency and effective mating frequency is most probably due to paternity skew within colonies. The study population proved to be genetically diverse and in Hardy-Weinberg equilibrium, suggesting random mating within the study area. No population substructure was observed, neither considering nuclear (global F _ST = 0.011 ± 0.003 SE) nor mitochondrial markers (mean Φ_ST = 0.008). Consequently, gene flow is obviously promoted by both sexes across the range investigated here. Thus, multiple mating and long-distance dispersal appear to be two interconnected behavioural mechanisms to create and maintain genetic diversity in At. colombica. The advantages of this system are partly offset by paternity skew and the non-zero relatedness among colony fathers found in the study population. Received 18 March 2008; revised 14 July 2008; accepted 18 July 2008.     

Polyandry in the wild: temporal changes in female mating frequency and sperm competition intensity in natural populations of the tettigoniid Requena verticalis

Empirical tests of sexual selection theory generally utilize model systems under laboratory settings, and extend conclusions to evolutionary processes occurring in nature. The biological significance of laboratory findings will depend largely on the mating rates of females and patterns of paternity in natural populations, information on which is generally lacking. Here we use microsatellite markers to provide rare estimates of female mating rates and patterns of parentage in a species of tettigoniid, Requena verticalis, which has been used extensively to test theory on the evolution of male parental investment and its influence on the direction of sexual selection. We found that although the number of males having a genetic representation in the female's sperm stores was higher for females collected late in the breeding season than those collected early in the season, overall the female mating rate was lower than that expected from laboratory observations. Analysis of parentage of offspring produced by females at the end of the breeding season revealed that all males represented in the sperm stores fathered offspring, although paternity was biased away from that expected from random sperm utilization. The data show that the complete first male sperm precedence documented in laboratory studies of this species does not persist in natural populations. Our data provide a solid underpinning for conclusions drawn from laboratory studies of this species.     

Queen polymorphism in the North American harvester ant,Ephebomyrmex imberbiculus

Summary In the North American harvester antEphebomyrmex imberbiculus, in addition to dealate queens, wingless female reproductives occur that have greatly reduced ocelli and thoracic sutures (intermorphic queens). Both queen types are equivalent in function, and do not differ in ovarian morphology. Colonies may contain several inseminated and egg-laying intermorphic queens. We discuss queen polymorphism in respect to the biology of this desert-dwelling species.     

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.     

An experimental study of bamboo ants in western Amazonia

Numerous ant taxa naturally inhabit stems of live and dead Guadua bamboo (Bambusoidea, Poaceae) in western Amazonia. In an experiment at the onset of the wet season in Peru’s Manu National Park, we augmented potential nest sites in stems of live bamboo, dead bamboo and dead ca?a brava (Gynerium sagittatum, another woody grass) at five stations within each of ten bamboo patches and ten control areas outside those patches. Each experimental stem possessed three vacant and available internodes, pre-drilled with, respectively, large, small and linear holes, mimicking the range of forms of surveyed natural entrances. After 24 days, approximately 13% of 798 available internodes had been colonized, the majority by fragments of existing colonies. Ignoring entrance type, which did not affect colonization for any species or species group, and censoring non-independent internodes of the same stem, we used individual stems as independent sample units in other tests. One specialist in live bamboo (Camponotus longipilis), and a likely specialist in dead bamboo (Camponotus depressus), were identified based on overrepresentation in bamboo habitat and disproportionate occurrence in live or dead bamboo stems. A third species, Camponotus (Pseudocolobopsis sp.) was more abundant in bamboo areas but colonized both dead bamboo and dead ca?a. Relatively high abundance of standing dead stems in Guadua forests may account for the presence of a dead stem specialist. The experiment missed detecting specialization in one live culm specialist (Camponotus mirabilis), likely due to its failure to simulate conditions required for the species’ unique modes of colony establishment and spread into new culms. Most opportunistic stem nesters colonized dead bamboo at significantly greater rates than dead ca?a, but were either equally well represented in bamboo and control areas, or underrepresented in bamboo habitat. Given low colonization rates overall, underrepresentation in bamboo cannot be attributed to competition from bamboo specialists for nesting space. Rather, it may be due to combined effects of seasonal flooding of bamboo habitat, and greater importance of food limitation, relative to nest site limitation, in that habitat. Received 9 May 2005; revised 25 August 2005; accepted 29 August 2005.     

Determinants of intracolonial relatedness in Pogonomyrmex rugosus (Hymenoptera; Formicidae): mating frequency and brood raids

The genus Pogonomyrmex is one of three ant genera with an effective mating frequency (me) > 2.0. We developed microsatellites to determine me for P. rugosus because mating frequency of P. rugosus was known only from observational data which do not allow an estimate of me. We genotyped 474 workers from 20 colonies for two microsatellite loci. Observed mating frequencies ranged from 3 to 12 and me for P. rugosus was 4.71. Observed patriline frequencies were significantly different from the expected patriline frequencies generated with a simulated data set under the assumption of equal patriline representation. The available mating frequency data and phylogenetic information of the genus Pogonomyrmex suggest that multiple mating is the ancestral state in the North American Pogonomyrmex sensu stricto. Established P. rugosus colonies raid and destroy smaller conspecific colonies. During these raids ant workers were observed carrying pupae and larvae from the raided colony into the nest of the raiding colony. However, it was not clear whether raided brood emerged in the raiding colony and were subsequently recruited into the work force (intraspecific slavery) or were used as food (predation). Our analyses indicate 6 of 14 field colonies contained foreign P. rugosus workers (43%). The range of the intracolonial frequency of foreign workers collected directly from the nest entrance was between 4 and 28%.