Differential learning and memory by co-occurring ant species

Foragers of the antsMessor pergandei andPogonomyrmex rugosus experience differing levels of variability in the distribution of seeds they harvest due to species-specific differences in foraging behavior.Messor pergandei foragers experience more variable seed distributions and densities, learn to recognize a novel seed faster but forget this information faster thanP. rugosus, which experiences more constant seed distributions even in the same habitat. Rate of learning to recognize a novel seed species was negatively associated with measures of seed species diversity for both ants.Messor pergandei foragers respond to variation in seed density by varying number of seeds handled per seed harvested, whileP. rugosus foragers do not. Memory of a novel seed exceeds forager longevity, due perhaps to use of seed caches as a type of information center.     

Dietary similarity and foraging range of two seed-harvester ants during resource fluctuations

Summary Diets of desert seed-harvester ants Veromessor pergandei and Pogonomyrmex rugosus were studied for 3 years at two habitats where they are common and sympatric. Diets of the two species were similar, consisting mainly (87% of 23,913 seeds) of three annual plant species (Schismus arabicus, Plantago insularis, and Pectocarya recurvata). Diets converged following a drought in Winter/Spring 1984 which reduced seed production during this time. Foraging range of P. rugosus almost doubled following the drought while foraging range of V. pergandei remained constant. Neither ant species move their nests once established rendering them effectively sessile granivores. This limits the dietary options of a given colony and may explain interspecific dietary convergence.     

Soil texture as an influence on the distribution of the desert seed-harvester ants Pogonomyrmex rugosus and Messor pergandei

Summary Pogonomyrmex rugosus and Messor pergandei are ecologically similar species of desert seed-harvester ants that coexist in numerous areas throughout the Sonoran and Mohave Deserts. However, these two species also commonly segregate along physical gradients, with each species predominating in areas that differ in soil texture and/or topographic relief. Along gradients that included bajada and alluvial flat habitats, P. rugosus occurred alone in coarse-textured soils near mountains, while M. pergandei occurred alone in finer-textured soils further away. Conversely, along a vegetation gradient that included creosote bush and saltbush habitats, P. rugosus occurred alone in finer-textured soils than those occupied by either M. pergandei alone or both species in coexistence. However, in both situations clay content was significantly higher in areas occupied by P. rugosus alone, and at the latter site clay content was correlated with relative abundance of each species. Moreover, local distribution pattern of these two species may be related to the effects of clay on water retention, with retention being highest in areas occupied by P. rugosus alone. Differences in reproductive ecology may also affect these patterns as P. rugosus reproductive flights follow summer monsoon rains, while those of M. pergandei occur during the milder winter and spring.     

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.     

Indirect effects of granivory by harvester ants: plant species composition and reproductive increase near ant nests

Summary Of 36 plant species surveyed, 6 were significantly associated with nests of the desert seed-harvester ant Veromessor pergandei or Pogonomyrmex rugosus; two other plant species were significantly absent from ant nests. Seeds of two common desert annuals, Schismus arabicus and Plantago insularis, realize a 15.6 and 6.5 fold increase (respectively) in number of fruits or seeds produced per plant growing in ant nest refuse piles compared to nearby controls. Mass of individual S. arabicus seed produced by plants growing in refuse piles also increased significantly. Schismus arabicus, P. insularis and other plants associated with ant nests do not have seeds with obvious appendages attractive to ants. Dispersal and reproductive increase of such seeds may represent a relatively primitive form of ant-plant dispersal devoid of seed morphological specializations. Alternatively, evolution of specialized seed structures for dispersal may be precluded by the assemblage of North American seed-harvester ants whose workers are significantly larger than those ants normally associated with elaiosome-attached seed dispersal. Large worker size may permit consumption of elaiosome and seed.     

Foraging behaviour of harvesting ants determines seed removal and dispersal

Harvesting ants can affect the regeneration of plants through at least two different processes: seed removal and seed dispersal. We analyse the role of different foraging strategies of ants on patterns of seed removal and dispersal by three Messor species with considerable differences in their foraging systems. Messor capitatus workers rarely leave the nest in well-formed columns, while the other two species form foraging trails, with M. bouvieri forming temporary trails and M. barbarus foraging on a stable system of permanent foraging trails. Overall seed intake of M. capitatus colonies is considerably less than that of the two group-foraging species. There are also differences in the size of seeds collected: M. barbarus and M. capitatus harvest similar amounts of large and small seeds, while M. bouvieri harvests small seeds more intensely than large ones, due to the smaller size of the worker caste. The three Messor species differ in the percent of seed dropping of the different seed type and in the seed dispersal distance. Moreover, M. bouvieri and M. capitatus redistributed dropped seeds preferentially in bare soil and low sparse vegetation habitats, while M. barbarus redistributed seeds mainly in the high vegetation habitat. These results show that the foraging systems of these harvesting ants determine different patterns of seed removal and dispersal and, thus, affect the abundance and redistribution of seeds in the area.     

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

Harvester ants usually go through temporal fluctuations in environmental seed abundance and composition which could influence their behaviour and ecology. The aim of this study was to evaluate how these fluctuations influence the diet of Pogonomyrmex rastratus, P. pronotalis and P. inermis (Hymenoptera, Formicidae) in the central Monte desert during three consecutive growing seasons. Although seeds were the main item in the diet, these ants turned more generalist when seed abundance of the most consumed species (grasses Aristida spp., Trichloris crinita, Pappophorum spp., Digitaria californica and Stipa ichu) was low. Accordingly, diversity of items in the diet decreased with seed abundance in a logarithmical fashion, showing higher foraging efficiency for seeds at higher seed abundance. Seed diversity, however, was not related to seed abundance as ants always included several species in their diet, with alternating prevalence. The proportion of the most consumed species increased logarithmically in the diet of P. rastratus and P. pronotalis along with their abundance in the environment probably as a consequence of diet switching (from forb and shrub seeds to grass seeds) and by an increase in foraging efficiency at higher seed densities. In contrast, foraging activity of P. inermis was very low at low seed abundance and its diet included only the five grasses. Among the most consumed species, proportion in the diet was not associated with relative abundance in the environment. Aristida spp., Pappophorum spp. and D. californica were overall highly selected. However, the flexibility in the diet of P. pronotalis and P. rastratus and the low foraging activity of P. inermis during periods of low resource abundance could attenuate potential top‐down effects in the central Monte desert. This study shows that bottom‐up effects are important in ant‐seed interactions and should be considered when predicting and evaluating ants' effects on seed resources.     

Is foraging in the desert ant,Messor pergandei (Hymenoptera: Formiceidae), limited by water?

Abstract.

• 1 The seed-harvesting ant Messor (Veromessor) prrgandei (Mayr) is a common inhabitant of southwestern deserts of the U.S.A. Foragers vary in size from less than 1 mg to more than 10 mg in body mass and may travel over 80 m on a single foraging trip. Their small size, long foraging range, and hot, arid habitat suggest that water stress may limit foraging activity. We examined intercolony and interindividual variation in water loss of M.pergandei foragers under several different situations in the field.
• 2 Colonies differed significantly in minimum critical water content (W_c) of individual foragers (water content below which foragers are incapable of normal locomotion). In one colony small workers had disproportionately higher W_c than larger workers; in the other colony W_c was isometric with body size.
• 3 Groups of workers confined in the field approached W_c only after normal foraging stopped and substrate temperatures exceeded 45°C, while water content of individual foragers did not approach the W_c during normal foraging periods. Moreover, seed load and distance travelled did not negatively affect forager water content, as measured on return to the nest: indeed, our results suggest that forager hydration level may influence load selection and/or foraging distance. We conclude that, under normal circumstances, foraging in M.pergandei is not water-limited.

     

Ants climb plants because they cannot swim: ant presence on flowers during the flood season reduces the frequency of floral visitors

1. Seasonal changes in environments may not only affect habitat connectivity but may also affect its use by species and their interactions. Thus, during the flood season, ants are forced to develop survival strategies such as vertical plant migration.
2. According to this, it has been hypothesized that the presence of ants may directly affect plant-pollinator interactions.
3. Thus, we asked the following questions: (i) Are floral visitors of Hyptis brevipes expelled due to ant presence on inflorescences during the flood period? (ii) Is the ant effect mediated by the abundance of ants foraging on inflorescences? And, (iii) Does flower abundance predict the abundance of floral visits and ants?
4. We experimentally sampled 59 H. brevipes plants with and without ants during the flooded season, and observed no differences in flower abundance between ant treatments.
5. The probability of detaining floral visitors on H. brevipes increased with ant abundance and exceeded 50% possible repellency, but the probability of visitor deterrence was not related to flower abundance. Furthermore, the abundance of flowers did not predict the number of ants on H. brevipes individuals or the frequency of floral visits.
6. Consequently, ant repelling effects are pronounced when there are more ants foraging on plants. However, the ant repelling effect can be mitigated when plants flourish all year-round and exhibit higher concentrations of flowers in the dry months. Additionally, the different sexual functions of plants may present specific responses due to the explosive pollination mechanism associated with ant effects.

     

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.     

Influence of seed size on feeding preferences and diet composition of three sympatric harvester ants in the central Monte Desert,Argentina

Selective seed consumption by harvester ants may affect seed abundance and composition and, ultimately, plant communities. We evaluated the influence of seed size on preferences and diet of Pogonomyrmex mendozanus, P. rastratus, and P. inermis in the central Monte Desert, Argentina. In choice experiments with Pappophorum spp. seeds of different sizes, P. mendozanus and P. rastratus preferred large seeds, maximizing energy reward. P. inermis showed a less-marked preference for large seeds, which was probably due to morphological constraints imposed by its small body size. Under natural conditions, none of the three species selected larger Pappophorum spp. seeds probably because of high travel and handling costs. Seeds of intermediate size predominated in the diet of the three species but a slight size match was detected as P. mendozanus carried larger seeds than P. rastratus and this than P. inermis, matching body-size differences. Thus, ants probably maximize energy reward but face morphological restrictions and higher costs when carrying and holding large seeds. While seeds of intermediate size are the most vulnerable ones to ant predation, small seeds are favored, as they are abundant in the soil seed bank and lowly predated.     

Foraging,food selection and worker size in the seed harvesting ant <Emphasis Type="Italic">Messor bouvieri</Emphasis>

Interactions among granivores and seeds depend on the foraging behaviour and morphology of the granivores and on the attributes and availability of seeds. We investigated seed selection by the seed harvesting ant Messor bouvieri in three adjacent plant communities in Spain by relating the harvested seeds to those in the seed rain. Preference was positively correlated with seed size and abundance which accounted for 43% and 20% of the variance respectively. Contrary to predictions of central place foraging theory, the size of seeds harvested did not increase with distance from the nest. Inclusion of a less-preferred item in the diet was more strongly related to the abundance of more-preferred items (60% of the variance) than the abundance of the less-preferred item (14% of the variance). Worker size accounted for 20–30% of the variance in the size of harvested seeds, although small workers did not appear to be constrained by load size for the range of seeds available. The body size of ants was significantly larger in the community with the greatest proportion of large seeds, although this was not due to their ability to carry larger loads or due to the greater force required to crush these seeds. The strong preference of M. bouvieri for large seeds may have important consequences for the plant communities in which they forage. Received: 30 November 1999 / Accepted: 14 April 2000     

Does the morphology of animal foraging pits influence secondary seed dispersal by ants?

Secondary seed dispersal by ants (myrmecochory) is an important process in semi‐arid environments where seeds are transported from the soil surface to an ant nest. Microsites from which ants often remove seeds are the small pits and depressions made by native and exotic animals that forage in the soil. Previous studies have demonstrated greater seed retention in the pits of native than exotic animals, but little is known about how biotic factors such as secondary seed dispersal by ants affect seed removal and therefore retention in these foraging pits. We used an experimental approach to examine how the morphology of burrowing bettong (Bettongia lesueur), greater bilby (Macrotis lagotis), short‐beaked echidna (Tachyglossus aculeatus) and European rabbit (Oryctolagus cuniculus) foraging pits and ant body size influenced ant locomotion and seed removal from pits along an aridity gradient. Ants took 3.7‐times longer to emerge from echidna pits (19.6 s) and six‐times longer to emerge from bettong pits (30.5 s) than from rabbit pits (5.2 s), resulting in lower seed removal from bettong pits than other pit types. Fewer seeds were removed from pits when cages were used to exclude large body‐sized (>2 mm) ants. Few seeds were removed from the pits or surface up to aridity values of 0.5 (humid and dry sub‐humid), but removal increased rapidly in semi‐arid and arid zones. Our study demonstrates that mammal foraging pit morphology significantly affects ant locomotion, the ability of ants to retrieve seeds, and therefore the likelihood that seeds will be retained within foraging pits.     

Seed removal in a tropical North American desert: an evaluation of pre‐ and post‐dispersal seed removal in Stenocereus stellatus

• To determine seed removal influence on seed populations, we need to quantify pre‐ and post‐dispersal seed removal. Several studies have quantified seed removal in temperate American deserts, but few studies have been performed in tropical deserts. These studies have only quantified pre‐ or post‐dispersal seed removal, thus underestimating the influence of seed removal. We evaluated pre‐ and post‐dispersal seed removal in the columnar cactus Stenocereus stellatus in a Mexican tropical desert.
• We performed selective exclosure experiments to estimate percentage of seeds removed by ants, birds and rodents during the pre‐ and post‐dispersal phases. We also conducted field samplings to estimate abundance of the most common seed removers.
• Birds (10–28%) removed a higher percentage of seeds than ants (2%) and rodents (1–4%) during pre‐dispersal seed removal. Melanerpes hypopolius was probably the main bird removing seeds from fruits. Ants (62–64%) removed a higher percentage of seeds than birds (34–38%) and rodents (16–30%) during post‐dispersal seed removal. Pogonomyrmex barbatus was probably the main ant removing seeds from soil.
• Birds and ants are the main pre‐ and post‐dispersal seed removers in S. stellatus, respectively. Further studies in other S. stellatus populations and plants with different life forms and fruit types will contribute to evaluate seed removal in tropical American deserts.

     

Endophytic fungi alter foraging and dispersal by desert seed-harvesting ants

Endophytic fungi are thought to interact mutualistically with host plants by producing alkaloid metabolites that deter herbivory. Since such fungi are transmitted via seed in some grasses, the presence of endophytes may also protect plants from seed predators. We conducted seed choice experiments for two dominant seed harvesting ants, Pogonomyrmex rugosus in the Sonoran desert and Pogonomyrmex occidentalis at a higher elevation, riparian zone in Arizona, USA. Non-infected fescue (Festuca arundinacea) seeds and seeds infected with the endophytic fungus, Acremonium coenophialum, were presented to ant colonies in three different populations. Infected seeds were harvested less frequently than non-infected seed for the two populations of Pogonomyrmex rugosus but not for the population of Pogonomyrmex occidentalis. We also a conducted seed dispersal experiment for one population of Pogonomyrmex rugosus. Of the seeds that were harvested, most of the colonies discarded more infected seeds into refuse piles than expected by chance. Seeds discarded into refuse piles have greater germination success than surrounding areas. The most important interaction of endophytes and grasses may be deterrence of seed predation and enhancing the probability of germinating in favorable sites, since these processes directly increase plant fitness.     

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.     

Effects of season and population size on pollination and reproductive output in a Mediterranean shrub

• Pollinator guilds may change throughout extended flowering periods, affecting plant reproductive output, especially in seasonal climates. We hypothesised a seasonal shift in pollinator guild and an autumn reduction in pollinator abundance, especially in small and sparse populations.
• We recorded pollinator identity, abundance and behaviour in relation to flower density from plant to population throughout the extended flowering of Ononis tridentata. We evaluated female reproductive output by recording pollination success and pre‐dispersal seed predation in eight populations of contrasting size and density. Offspring quality was also characterised through seed weight and germination.
• A diverse guild of insects visited O. tridentata in spring, while only Apis mellifera was observed in autumn. Visitation frequency did not vary seasonally, but the number of flowers per foraging bout was lower, and seeds were heavier and had a higher germination rate in autumn. Plant and neighbourhood flowering display were not related to pollinator visitation frequency or behaviour. However, the rate of fertilised ovules, seed set and autumn flowering display size were positively related to population density.
• The maintenance of pollination in autumn enhances the reproductive performance of O. tridentata due to higher quality of autumn seed, and to a large reduction in seed predator pressure. We also suggest that observed changes in pollinator behaviour could be one of the processes behind seasonal variation in seed performance, since geitonogamous crosses were less likely to occur in autumn.

     

Dropping rates of elaiosome-bearing seeds during transport by ants (Formica polyctena Foerst.): Implications for distance dispersal

In a deciduous forest, foraging ants collect elaiosome-bearing seeds and carry them to their nests. Some of the seeds reach the nest and are concentrated there. Others may be dropped by ants during transport. The dropped seeds enter the soil seed pool. However, they might be repeatedly removed by other ant individuals and carried again in the direction of the nest. Rates of seed dropping and repeated removals must be known to evaluate the effect of ant workers on dispersal distance of seeds. The rate of seed dropping is predicted to depend on size of seeds and of elaiosomes, both of which vary among plant species, and on the size of the ant workers. Mark-recapture experiments were used to evaluate dropping rates of seeds of five myrmecochorous and diplochorous plants (Chelidonium majus L., Asarum europaeum L., Viola matutina Klok., V. mirabilis L., V. hirta L.) during their transport by the ant Formica polyctena Foerst. In the series of species A. europaeum – V. hirta – V. mirabilis – Ch. majus – V. matutina, the dropping rate increased. Small workers dropped seeds of A. europaeum more often than did large ones, while seeds of V. hirta were dropped by ants of different size classes with the same frequency. Across species, dropping rates of seeds were negatively correlated with the rate at which ants removed them from the depot. The number of seeds which reach the nests is the other important parameter of seed dispersal. This parameter depends on dropping rates: seeds with lower dropping rates have higher chances of being deposited in nests. Diplochores usually produce many small seeds, which are characterised by low removal rates and high dropping rates during transport by ants. Obligate myrmecochores produce rather few large seeds, which have high removal rates and low dropping rates. To analyse the significance of seed dropping in the dispersal distance of seeds, a computer simulation based on two factors [(i) seed number produced by a plant; (ii) dropping rate of seeds] is proposed.     

Foraging behavior and morphology: seed selection in the harvester ant genus, Pogonomyrmex

Optimally foraging animals can be behaviorally or morphologically adapted to reduce the energetic and time costs of foraging. We studied the foraging behavior and morphology of three seed harvester ant species, Pogonomyrmex barbatus, P. desertorum, and P. occidentalis, to determine the importance of behavioral strategies and morphological features associated with load carriage in reducing the costs of foraging. We found that none of five morphological features we measured had a significant impact on seed selection. Also, body size did not influence running speed, an important variable in time costs of foraging. Temperature had the largest effect on running speed in these species. Our results show that these species have foraging strategies which minimize the time costs of traveling with seeds. We also describe a pattern where the running speed in individual-foraging species is less affected by increasing seed size than in trunk-trail foragers, when temperature and body mass are held constant. These results support previous work which showed that time costs are most important in seed selection for Pogonomyrmex, and suggest that central place foraging theory may need to accommodate variation in foraging strategy to more accurately predict optimal seed size selection in harvester ants. Received: 16 June 1997 / Accepted: 15 December 1997     

Germination niche breadth varies inconsistently among three Asclepias congeners along a latitudinal gradient

• Species responses to climate change will be primarily driven by their environmental tolerance range, or niche breadth, with the expectation that broad niches will increase resilience. Niche breadth is expected to be larger in more heterogeneous environments and moderated by life history. Niche breadth also varies across life stages. Therefore, the life stage with the narrowest niche may serve as the best predictor of climatic vulnerability. To investigate the relationship between niche breadth, climate and life stage we identify germination niche breadth for dormant and non‐dormant seeds in multiple populations of three milkweed (Asclepias) species.
• Complementary trials evaluated germination under conditions simulating historic and predicted future climate by varying cold–moist stratification temperature, length and incubation temperature. Germination niche breadth was derived from germination evenness across treatments (Levins B_n), with stratified seeds considered less dormant than non‐stratified seeds.
• Germination response varies significantly among species, populations and treatments. Cold–moist stratification ≥4 weeks (1–3 °C) followed by incubation at 25/15 °C+ achieves peak germination for most populations. Germination niche breadth significantly expands following stratification and interacts significantly with latitude of origin. Interestingly, two species display a positive relationship between niche breadth and latitude, while the third presents a concave quadratic relationship.
• Germination niche breadth significantly varies by species, latitude and population, suggesting an interaction between source climate, life history and site‐specific factors. Results contribute to our understanding of inter‐ and intraspecific variation in germination, underscore the role of dormancy in germination niche breadth, and have implications for prioritising and conserving species under climate change.