Coffee farm diversity and landscape features influence density of colonies of Atta cephalotes (Hymenoptera: Formicidae)

The density of colonies of leaf-cutting ants, Atta cephalotes L. (Hymenoptera: Formicidae), was measured and compared among coffee (Coffea arabica L.) plantations in five management categories: monoculture conventional, diversified conventional, diversified organic, highly diversified conventional, and highly diversified organic. Twenty-four small farms (<4 ha) in Turrialba, Costa Rica, were included in this study. Within-farm and off-farm (landscape) variables were measured and tested for their relationship with A. cephalotes colony density. Total ant colony density (colonies per ha) and density of new colonies shortly after a nuptial flight were significantly greater in the coffee monoculture conventional system, compared with all other systems. Total ant colony density and density of new colonies were inversely related to percentage of shade within the farms. Within farms, colony density was greater near edges adjacent to riparian forest than those adjacent to nonforested land. Regardless of edge type, plots closer to the edge (0-30 m) had greater colony density than those furthest from the edge. At the landscape scale, density of new colonies was positively related to fallow land use coverage within a 2,000-m buffer radius and to forest coverage within a 500-m radius. Results indicate that coffee farm management practices and landscape level factors can affect A. cephalotes colony densities. Understanding such practices and factors could assist in the development of better management methods of these injurious insects in coffee farms. Increased diversification in coffee farms, possibly due to the greater shade associated with it, may reduce colonization by the ants, which are considered forest gap specialists.     

Cutting More from Cut Forests: Edge Effects on Foraging and Herbivory of Leaf-Cutting Ants in Brazil

Edge-mediated changes in species composition are known to result in modified species interactions. Because of the crucial trophic position of herbivores and their far-reaching impact on plant communities, it is important to understand how edge influences herbivory. In the present paper, we investigated whether and how leaf-cutting ant foraging is altered in the forest edge, as this habitat is characterized by an increased proportion of pioneer species. We assessed basic foraging data as well as the herbivory rate ( i.e. , the proportion of the leaf material harvested by a colony in relation to the available leaf area in the foraging area) of Atta cephalotes colonies at the edge versus interior sites of a large remnant of the Atlantic forest in Northeast Brazil. Our results indicated clear edge effects on leaf-cutting ants: equally sized A. cephalotes colonies located at the forest edge removed about twice as much leaf area from their foraging grounds than interior colonies (14.3 vs. 7.8%/col/yr). This greater colony-level impact within the forest edge zone was a consequence of markedly reduced foraging areas (0.9 vs. 2.3 ha/col/yr) and moderately lower leaf area index in this habitat, whereas harvest rates were the same. Our results suggest that forest edges induce increased leaf-cutting ant herbivory, probably via the release of resource limitation. Together with the increase of leaf-cutting ant populations along forest edges, this may amplify environmental changes induced by habitat fragmentation.     

A critical analysis of parameter adaptation in ant colony optimization

Applying parameter adaptation means operating on parameters of an algorithm while it is tackling an instance. For ant colony optimization, several parameter adaptation methods have been proposed. In the literature, these methods have been shown to improve the quality of the results achieved in some particular contexts. In particular, they proved to be successful when applied to novel ant colony optimization algorithms for tackling problems that are not a classical testbed for optimization algorithms. In this paper, we show that the adaptation methods proposed so far do not improve, and often even worsen the performance when applied to high performing ant colony optimization algorithms for some classical combinatorial optimization problems.     

Invasive alien plants affect grassland ant communities, colony size and foraging behaviour

Ants are dominant members of many terrestrial ecosystems and are regarded as indicators of environmental changes. However, little is known about the effects of invasive alien plants on ant populations, particularly as regards the density, spatial distribution and size of ant colonies, as well as their foraging behaviour. We addressed these questions in a study of grassland ant communities on five grasslands invaded by alien goldenrods (Solidago sp.) and on five non-invaded grasslands without this plant. In each grassland, seven 100 m² plots were selected and the ant colonies counted. Ant species richness and colony density was lower in the plots on the invaded grasslands. Moreover, both of these traits were higher in the plots near the grassland edge and with a higher number of plant species in the grasslands invaded by goldenrods but not in the non-invaded ones. On average, ant colony size was lower on the invaded grasslands than the non-invaded ones. Also, ant workers travelled for longer distances to collect food items in the invaded areas than they did in the non-invaded ones, even after the experimental removal of some ant colonies in order to exclude the effect of higher colony density in the latter. Our results indicate that invasive alien goldenrods have a profound negative effect on grassland ant communities which may lead to a cascade effect on the whole grassland ecosystem through modification of the interactions among species. The invasion diminishes a major index of the fitness of ants, which is a colony’s size, and probably leads to increased foraging effort of workers. This, in turn, may have important consequences for the division of labour and reproductive strategies within ant colonies.     

Ants impact the composition of the aquatic macroinvertebrate communities of a myrmecophytic tank bromeliad

In an inundated Mexican forest, 89 out of 92 myrmecophytic tank bromeliads (Aechmea bracteata) housed an associated ant colony: 13 sheltered Azteca serica, 43 Dolichoderus bispinosus, and 33 Neoponera villosa. Ant presence has a positive impact on the diversity of the aquatic macroinvertebrate communities (n = 30 bromeliads studied). A Principal Component Analysis (PCA) showed that the presence and the species of ant are not correlated to bromeliad size, quantity of water, number of wells, filtered organic matter or incident radiation. The PCA and a generalized linear model showed that the presence of Azteca serica differed from the presence of the other two ant species or no ants in its effects on the aquatic invertebrate community (more predators). Therefore, both ant presence and species of ant affect the composition of the aquatic macroinvertebrate communities in the tanks of A. bracteata, likely due to ant deposition of feces and other waste in these tanks.     

Dirt roads and fire breaks produce no edge effects on litter-dwelling arthropods in a tropical dry-forest: a case study

Edge effects threaten organisms and ecological processes in habitat remnants, but they have been poorly studied in non-humid forests such as cerradão, a tropical dry forest sometimes derived from fire-suppressed savanna in Brazil. The diverse ecosystem functions performed by arthropods may be disrupted by edge effects, and there is pressing need for more studies on this subject. We sampled fragments of cerradão facing either a road or fire breaks, assessing edge effects in: beta diversity and community composition of epigaeic (litter-dwelling) arthropod orders, ant species, and ant functional groups; ant species richness and diversity; leaf litter depth; and colony residence time of a predatory ground-dwelling ant, Odontomachus chelifer (Ponerinae). None of the variables measured differed between edge and interior of the sites sampled. Dry forests have high micro-climatic variations caused by discontinuities in the canopy cover and, as such, changes in abiotic variables in cerradão edges might not be as clear as those observed in tropical rainforests. Our study demonstrates that edge effects may not be so prevalent in cerradão facing roads or fire breaks, which possibly increases the chances of survival of a higher fraction of the original arthropod fauna compared to rainforest fragments.     

基于蚁群遗传算法的DNA序列比对方法

蚁群遗传算法是在蚁群算法的基础上用遗传算法对其参数进行优化而产生的一种改进算法。把蚁群遗传算法应用于DNA序列比对上,结果表明这种新的序列比对算法是非常有效的。     

Characterizing critical rules at the 'edge of chaos'

This paper aims to provide a deeper understanding of the 'edge of chaos' phenomenon. This is an important concept in some strands of work on complex systems, but exactly what it means remains unclear. Broadly, it may be seen as a region between rigid order and chaos. In cellular automata which show the edge of chaos phenomenon, this is indicated by the occurrence of smooth transitions in certain variables, such as the average Shannonian entropy, where generally one finds sudden jumps. The abrupt jump is between an ordered realm and a chaotic one, so the smoothness of a transition holds out the promise of the existence of a realm between order and chaos. Since living systems may also be described as lying between order and chaos, then the edge of chaos may be the key to understanding the form of organization which characterizes life. The smooth transitions in 2-D cellular automata were therefore investigated in detail to see whether it is possible to characterize the cellular automata rules which give rise to them. It was found that the transitions were smoother the less that there was a marked jump in the number of rule table entries applied. But, the bare fact of this correspondence does not reveal the cause of the smoothness. Further work was therefore carried out to try to understand more about what led to the smooth behaviour. However in two further studies in which 'smooth rules' were slightly altered and in which the dominant parts of a 'smooth rule' were imposed on other rules, it was found that the rules which led to a smooth transition were very sensitive to small changes. A change of less than 0.05% of the rule table entries of a 'smooth rule' was more than likely to lead to an abrupt rather than smooth transition. This suggests that it may prove difficult to understand what it is about the rules which leads to the edge of chaos.     

Estimation-based ant colony optimization and local search for the probabilistic traveling salesman problem

The use of ant colony optimization for solving stochastic optimization problems has received a significant amount of attention in recent years. In this paper, we present a study of enhanced ant colony optimization algorithms for tackling a stochastic optimization problem, the probabilistic traveling salesman problem. In particular, we propose an empirical estimation approach to evaluate the cost of the solutions constructed by the ants. Moreover, we use a recent estimation-based iterative improvement algorithm as a local search. Experimental results on a large number of problem instances show that the proposed ant colony optimization algorithms outperform the current best algorithm tailored to solve the given problem, which also happened to be an ant colony optimization algorithm. As a consequence, we have obtained a new state-of-the-art ant colony optimization algorithm for the probabilistic traveling salesman problem.     

Field experiments show contradictory short‐ and long‐term myrmecochorous plant impacts on seed‐dispersing ants

1. Some interactions previously described as mutualistic were revealed to be commensal or parasitic in subsequent investigations. Ant‐mediated seed dispersal has been described as a mutualism for more than a century; however, recent research suggests that it may be commensal or parasitic. Plants demonstrably benefit from ant‐mediated seed dispersal, although there is little evidence available to demonstrate that the interaction benefits long‐term ant fitness. 2. Field experiments were conducted in temperate North America focused on a key seed‐dispersing ant. All herbaceous plants were removed from a forest understorey for 13 years, and supplemented ant colonies with large elaiosome‐bearing seeds aiming to examine potential long‐ and short‐term myrmecochorous plant benefits for the ants. 3. If elaiosome‐bearing seeds benefit ants, suggesting a mutualistic relationship, it is expected that there would be greater worker and/or alate abundance and greater fat reserves (colony lipid content) with seed supplementation (short‐term) and in areas with high understorey herb abundance. 4. Short‐term seed supplementation of ant colonies did not result in an increase with respect to numbers or fat stores, although it did prompt the production of colony sexuals, which is a potential fitness benefit. In the long term, however, there was no positive effect on the ants and, instead, there were negative effects because the removal of elaiosome‐bearing plants corresponded with greater colony health. 5. The data obtained in the present study suggest that the ant–plant interaction ranged from occasionally beneficial to neutral to overall negative for the ant partner. Such results did not support considering the interaction as a mutualism. Collectively, the data suggest the need to reconsider the nature of the relationship between these ants and plants.     

Why do ants shift their foraging from extrafloral nectar to aphid honeydew?

When aphids parasitize plants with extrafloral nectaries (EFNs) and aphid colony size is small, ants frequently use EFNs but hardly tend aphids. However, as the aphid colony size increases, ants stop using EFNs and strengthen their associations with aphids. Although the shift in ant behavior is important for determining the dynamics of the ant–plant–aphid interaction, it is not known why this shift occurs. Here, we test two hypotheses to explain the mechanism responsible for this behavioral shift: (1) Extrafloral nectar secretion changes in response to aphid herbivory, or (2) plants do not change extrafloral nectar secretion, but the total reward to ants from aphids will exceed that from EFNs above a certain aphid colony size. To judge which mechanism is plausible, we investigated secretion patterns of extrafloral nectar produced by plants with and without aphids, compared the amount of sugar supplied by EFNs and aphids, and examined whether extrafloral nectar or honeydew was more attractive to ants. Our results show that there was no inducible extrafloral secretion in response to aphid herbivory, but the sugar concentration in extrafloral nectar was higher than in honeydew, and more ant workers were attracted to an artificial extrafloral nectar solution than to an artificial aphid honeydew solution. These results indicate that extrafloral nectar is a more attractive reward than aphid honeydew per unit volume. However, even an aphid colony containing only two individuals can supply a greater reward to ants than EFNs. This suggests that the ant behavioral shift may be explained by the second hypothesis.     

Variation in thermal tolerance of North American ants

Changing climates are predicted to alter the distribution of thermal niches. Small ectotherms such as ants may be particularly vulnerable to heat injury and death. We quantified the critical thermal maxima of 92 ant colonies representing 14 common temperate ant species. The mean CT_max for all measured ants was 47.8 °C (±0.27; range=40.2–51.2 °C), and within-colony variation was lower than among-colony variation. Critical thermal maxima differed among species and were negatively correlated with body size. Results of this study illustrate the importance of accounting for mass, among and within colony variation, and interspecific differences in diel activity patterns, which are often neglected in studies of ant thermal physiology.     

Theoretical analysis of two ACO approaches for the traveling salesman problem

Bioinspired algorithms, such as evolutionary algorithms and ant colony optimization, are widely used for different combinatorial optimization problems. These algorithms rely heavily on the use of randomness and are hard to understand from a theoretical point of view. This paper contributes to the theoretical analysis of ant colony optimization and studies this type of algorithm on one of the most prominent combinatorial optimization problems, namely the traveling salesperson problem (TSP). We present a new construction graph and show that it has a stronger local property than one commonly used for constructing solutions of the TSP. The rigorous runtime analysis for two ant colony optimization algorithms, based on these two construction procedures, shows that they lead to good approximation in expected polynomial time on random instances. Furthermore, we point out in which situations our algorithms get trapped in local optima and show where the use of the right amount of heuristic information is provably beneficial.     

The Relationship between Canopy Cover and Colony Size of the Wood Ant Formica lugubris - Implications for the Thermal Effects on a Keystone Ant Species

Climate change may affect ecosystems and biodiversity through the impacts of rising temperature on species’ body size. In terms of physiology and genetics, the colony is the unit of selection for ants so colony size can be considered the body size of a colony. For polydomous ant species, a colony is spread across several nests. This study aims to clarify how climate change may influence an ecologically significant ant species group by investigating thermal effects on wood ant colony size. The strong link between canopy cover and the local temperatures of wood ant’s nesting location provides a feasible approach for our study. Our results showed that nests were larger in shadier areas where the thermal environment was colder and more stable compared to open areas. Colonies (sum of nests in a polydomous colony) also tended to be larger in shadier areas than in open areas. In addition to temperature, our results supported that food resource availability may be an additional factor mediating the relationship between canopy cover and nest size. The effects of canopy cover on total colony size may act at the nest level because of the positive relationship between total colony size and mean nest size, rather than at the colony level due to lack of link between canopy cover and number of nests per colony. Causal relationships between the environment and the life-history characteristics may suggest possible future impacts of climate change on these species.     

Combining distances of ballistic and myrmecochorous seed dispersal in Adriana quadripartita (Euphorbiaceae)

The separate contributions of different vectors to net seed dispersal curves of diplochorous systems have rarely been characterised. In Australia, myrmecochory is a common seed dispersal syndrome and in the majority of such systems, seeds are initially dispersed ballistically. We measured ballistic and myrmecochorous seed dispersal distances in relation to canopies of Adriana quadripartita (Euphorbiaceae) and used a simulation model to estimate the net dispersal curve. We also compared seed removal rates and ant abundances under, and outside, plant canopies to examine how foraging patterns by ants may affect net dispersal.Overall ant abundance did not show a significant numerical response to seedfall; however, the abundance of the main seed dispersing ant, Rhytidoponera ‘metallica’ did. Despite this, seed removal rates did not differ significantly between canopy and open locations. Rhytidoponera ‘metallica’ account for 93% of observed seed dispersal events. On average, the ants dispersed seeds 1.54 m and in doing so, moved seed a mean radial distance of 0.76 m away from canopy edges. This contribution to net dispersal distance by ants is considerable since ballistic dispersal moved seeds a median distance of 7.5 cm. Our simulation model indicated that the combination of ballistic and ant seed dispersal is expected to result in seeds being transported a median net radial dispersal distance of 1.05 m from the canopy edge.Thus in this system, an important function of diplochory may simply be to move a higher proportion of seeds from under the canopy of parent plants than is possible by ballistic dispersal alone. This ‘dispersal-for-distance’ may result in reduced parent–offspring competition or may increase the probability that seeds reach rare safe sites for germination and recruitment.     

Edge effects decrease ant-derived benefits to seedlings in a neotropical savanna

Edge effects may lead to changes in mutualistic plant–animal interactions, such as seed dispersal, that are critical to plant regeneration. However, research into edge effects is neglected in the Brazilian cerrado, the largest neotropical savanna. We evaluated the consequences of edge effects in the cerrado for the regeneration of Erythroxylum pelleterianum (Erythroxylaceae), a shrub that benefits from seed dispersal by ants. We compared air temperature, relative humidity, and vapor pressure deficit, as well as the frequency and outcome of ant–diaspore interactions between cerrado edges and interiors. The inner portion of cerrado was likely to be moister than its borders, but seed production and germination did not differ between edge and interior of cerrado. Ants removed more seeds near fragment edges than at the interior. However, Myrmicinae ants dominated ant–fruit interactions at edges. These ants are likely to provide few benefits to the seeds. Seedlings of E. pelleterianum growing close to Ponerinae ant nests showed higher survival than seedlings growing away from nests in the interior of cerrado, but such effect disappeared near edges. Widespread seedling mortality due to a higher evaporative demand at edges may partially account for this effect. Furthermore, Ponerinae’s nests also showed a lower residence time near edges, decreasing possible benefits derived from ant colony activity such as nutrient enrichment and protection against insect herbivores. Edge effects could change the structure and dynamics of vegetation in cerrado fragments, due in part to the collapse of the mutualistic interactions demonstrated here.     

Effect of neighboring seeds associated with habitats on the seed removal of two coexisting myrmecochorous plants in central China

Although neighboring plants can influence animal-seed dispersal interactions, little is known about the effect of neighboring seeds and the influence of habitat on seed dispersal by ants. Here we investigated the influence of neighboring seeds on seed removal in two coexisting myrmecochorous species (Epimedium pubescens and Helleborus thibetanus) from temperate deciduous forests of Qinling Mountains, central China, by examining (1) the potential role of ants and rodents and (2) whether the neighboring seed effect differed between forest edge and interior. We found that, presence of the higher-ESMR (elaiosome: seed mass ratio) E. pubescens did not significantly affect seed removal of the lower-ESMR H. thibetanus. By contrast, the presence of H. thibetanus decreased the seed removal rates of E. pubescens, with only a significant effect in rodent exclusion (ant alone) rather than in both ant exclusion (rodent alone) and full access (rodent + ant). Moreover, we found that those effects were not significantly different between the forest edge and the interior, which may be attributed to a similar pattern of overall seed-dispersing ant abundance in the two habitats. This suggested that neighboring seeds could influence seed removal of the focal plant depending on the absence of rodents; when rodents were present, the interaction of rodents and ants rendered no influence of neighboring seeds on seed removal. Our results show that the neighborhood effect was regulated by both dispersers and predators, and this effect was not context-dependent at a small spatial scale. This study highlights the importance to understand the effect of shared seed-removing animals and habitat context to assess the neighboring seed effect on plant-animal interactions.     

Queen transport during ant colony emigration: a group-level adaptive behavior

Ant colonies emigrate frequently from one nest site to another. Emigrations, however, are dangerous, particularly for colonieswith a single queen. The queen is a "vital organ" of the colony,and emigrations expose her to grave peril. The optimal strategyfor a monogynous ant colony, therefore, should be that thequeen moves during the middle of the emigration so that sheis transferred swiftly from the protection of half of the colony in the old nest to the protection of the other half colony inthe new nest. In the ant Leptothorax albipennis, the queenis carried during colony emigration. We tested the null hypothesisthat the queen has a random position in the sequence of transportevents during an emigration. The result of 32 emigrations demonstrated,for the first time, that the transport serial number of thequeen [calculated relative to the total number of all transportevents (i.e., of brood and adult ants together), brood transportevents, or adult ant transport events] is not random and furthermoreoccurs in the middle of the transport sequence. This resultrepresents a colony strategy because we found that the relativetransport serial number of the queen was related neither to emigration distance nor to colony size. Transporting queensin the middle of emigrations is a strategy probably favoredby selection and is an aspect of colonies behaving as group-leveladaptive units.     

The relationship between the distribution of worker sizes and new worker production in the ant Formica neorufibarbis

While it is commonly assumed that variation in worker sizes within a single ant colony increases colony efficiency, there is little causal evidence of a link between worker size variation and colony performance. I tested whether the range of worker sizes within colonies of the ant species Formica neorufibarbis affected new worker production. Removing large workers from colonies lowered the rate of new worker production. A study of unmanipulated colonies indicated that colonies did not maintain a full range of worker sizes; mean worker head widths varied from 0.89-1.24 mm. Colonies naturally missing large workers did not have lower rates of worker production, suggesting that the relative size, not the absolute size, of workers within colonies was important. These are the first results to directly link the range of worker sizes to a component of colony fitness in a natural setting.     

Does the host plant affect the benefits from mutualisms? The invasive mealybug and ghost ant association

1. The benefits to trophobionte hemipterans are affected by the ant tending level, which is a widely accepted statement. The ant tending level is closely related to multiple factors. It is clear that the ant tending level can be affected by the temporal factor, age‐specific, the density of the hemipterans, and quantity and quality of honeydew produced by hemipterans. 2. Few studies of ant–hemipteran mutualisms have reported the patterns of host plants‐dependent effects, and whether host plants influence the ant tending level that is also unclear. As such, laboratory experiments were conducted to test whether the colony growth rate of an invasive mealybug Phenacoccus solenopsis Tinsley, parasitism of Aenasius bambawalei Hayat, an dominant parasitoid of P. solenopsis, are affected by tending by ghost ants (Tapinoma melanocephalum(Fabricius)], host plants (tomato and cotton), and interactions between the two factors. The difference in the ant tending level between the host plants was also determined. 3. The results showed that mealybug colony growth and parasitism were significantly affected by ant tending and host plant separately. There were significant interactions between the independent factors on the mealybug colony growth rate and percentage parasitism. These results suggest that benefits to mealybugs are host plant‐dependent.