Trail following,speed and fractal dimension of movement in a marine prosobranch,Littorina littorea,during a mating and a non-mating season

We quantitatively studied movement behaviour in the gastropod Littorina littorea in laboratory experiments during periods of their non-mating season (November 1992) and their mating season (April–May 1993). Snails were collected in 1992 and 1993 from one boulder shore on the north west coast of Sweden. In a comparison between the two seasons (one non-mating and one mating) we measured trail complexity of unsexed snails using fractal dimesion, the degree of mucus trail following (coincidence index of marker and tracker trails) and average movement speed of marker and tracker snails. We found no differences in fractal dimension and coincidence index of trails between the two seasons. Tracker snails moved, however, significantly faster than marker snails during both seasons. This could not be explained by trackers, when locomoting, using the mucus trail deposited by the marker to increase their speed, since there was no correlation between coincidence index and tracker speed. During the mating season we also conducted trail complexity, trail following and speed experiments comparing the behaviour of males and females. There was no difference between males and females in the fractal dimension of movement, nor was there any difference between the mean speed of male and female snails, although male marker snails tended to move faster than female marker snails. Males tracking other males, females tracking other females and females tracking males followed trails about equally long distances (i.e., coincidence indices did not differ). In contrast, males following female mucus trails showed a significantly higher degree of trail following than the other sex combinations. This new finding may suggest that females of L. littorea release pheromones in their mucus trails and that males are able to identify them.     

Different habitats, different habits? Response to foraging information in the parasitic wasp Venturia canescens

The parasitic wasp, Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae), has two reproductive modes, namely, thelytoky or arrhenotoky, and occurs in habitats with highly variable or relatively stable host abundances, respectively. Since information processing is costly, we expected that information indicating resource availability would be mainly used by the thelytokous wasps and less so by the arrhenotokous type. This idea was explored by two different approaches. In a study on patch-time allocation, we used females from ten populations and measured patch-residence times of individuals that visited multiple patches at different encounter rates. In a more detailed approach, thelytokous and arrhenotokous females from a single location were observed continuously while foraging, and all behaviors were recorded. Wasps of both reproductive modes (i.e., both habitat types) used information for the assessment of habitat quality. However, the way that the information was used differed between them. Whereas thelytokous females used foraging information to maximize their efficiency at high patch-encounter rates, arrhenotokous females merely reduced the number of offspring produced without changing patch times. The behavior of the arrhenotokous females should result in a spreading of offspring across the habitat and, thus, reduced sib-mating. The foraging strategy of these wasps might therefore be an adaptation to reduce costs associated with inbreeding.     

Division of foraging labor in ants can mediate demands for food and safety

Solitary foragers can balance demands for food and safety by varying their relative use of foraging patches and their level of vigilance. Here, we investigate whether colonies of the ant, Formica perpilosa, can balance these demands by dividing labor among workers. We show that foragers collecting nectar in vegetation near their nest are smaller than are those collecting nectar at sites away from the nest. We then use performance tests to show that smaller workers are more likely to succumb to attack from conspecifics but feed on nectar more efficiently than larger workers, suggesting a size-related trade-off between risk susceptibility and harvesting ability. Because foragers that travel away from the nest are probably more likely to encounter ants from neighboring colonies, this trade-off could explain the benefits of dividing foraging labor among workers. In a laboratory experiment, we show that contact with aggressive workers results in an increase in the mean size of recruits to a foraging site: this increase was not the result of more large recruits, but rather because fewer smaller ants traveled to the site. These results suggest that workers particularly susceptible to risk avoid dangerous sites, and suggest that variation in worker size can allow colonies to exploit profitably both hazardous and resource-poor patches.Communicated by L. Sundström     

Determinants of multiple central-place territory use in wild young-of-the-year Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

Patterns of space use provide key insights into how animals exploit local resources and are linked to both the fitness and distribution of individuals. We studied territory size, mobility, and foraging behavior of young-of-the-year Atlantic salmon Salmo salar in relation to several key environmental factors in Catamaran Brook, New Brunswick, Canada. The 50 study fish were all multiple central-place foragers (i.e., alternated among several sit-and-wait foraging stations) and showed great variability in territory size and the total distance traveled within the territories. Territory size increased with the mean distance traveled between consecutive foraging stations, the number of stations visited, and the mean foraging radius. Fish also varied greatly in how much of the total travel distance was associated with foraging at a station (14.8–91.8%) versus switching among stations (4.6–84.3%). As predicted, fish in slow-flowing waters, where drifting prey were scarce, used larger multiple central-place territories than individuals in faster, more productive waters. Interestingly, however, the most mobile fish did not inhabit slow-running waters as predicted but were found at intermediate (optimal) water current velocities. Hence, our study suggests that among some multiple central-place foragers, increased mobility may not only serve to increase prey encounter rate but may reflect an attempt to patrol territories in favorable habitats. Further studies are needed to determine the generality and the ultimate benefits of multiple central-place space use among stream-dwelling fish and other animals.     

Maximization of foraging efficiency and resource defense by group retrieval in the ant Formica schaufussi

Summary Workers of the ant Formica schaufussi forage as individuals and cooperate in groups to retrieve arthropod prey. In 2 sample years, group-transported prey were on average 6.8 and 4.7 times heavier than individually retrieved items, and the average loading ratios of groups were greater than the loading ratios of single foragers. Retrieval group size was adjusted to prey size, and prey transport velocity for individuals and groups tended to decrease with increasing prey weight. The efficiency of individual and group retrieval, estimated from calculations of the prey delivery rate to the nest (PDR) achieved by each foraging mode, varied as a function of prey size. Individual retrieval maximized PDR at a prey weight of 19.5 mg, and group transport maximized PDR at 190 mg. Although the PDR maxima of an individual in a group and a solitary forager were approximately equal, depending on prey size, group transport may maximize foraging efficiency. Group transport also decreased interference competition from sympatric ant species. Group-transported prey having a greater likelihood of successful retrieval were within the size range of prey that maximized foraging efficiency. Transport group size appeared to be more important in prey defense than in increasing prey transport velocity, suggesting an important role of group size in competitive ability.Offprint requests to: J.F.A. Traniello     

Oceanographic characteristics of areas used by Cory’s shearwaters during short and long foraging trips in the North Atlantic

Many breeding seabird species are central-place foragers and constrained to find productive prey patches within their foraging ranges. We assessed how different populations of a pelagic seabird species, the Cory’s shearwater Calonectris diomedea, breeding in oceanic and neritic conditions, cope with these constraints in the North Atlantic, during both incubation and chick-rearing periods. We analysed 237 foraging trips to study the movements and oceanographic characteristics of foraging habitats of seven different populations of Cory’s shearwaters. Generally, oceanic populations exhibited higher foraging effort, by travelling more time and to more distant areas, and larger home ranges and feeding areas, than the neritic population (i.e. breeding on an island within the Portuguese continental platform). On their short trips (i.e. ≤4 days), birds from the different populations fed mostly in shallower waters around the colony. During long trips (i.e. ≥5 days), feeding areas of both oceanic and neritic populations were characterized by high concentration values of chlorophyll-a, low sea-surface temperature and shallower habitats, with oceanic populations of the Azores exploiting areas north of the islands over known seamounts and frontal regions. Birds from other oceanic population (Selvagens) also exploited the African continental shelf system on their long trips. The home ranges of the different populations overlapped widely, but there was a general spatial segregation in terms of the core feeding areas at the population level. Core feeding areas and areas of foraging overlap between different populations should be important to inform conservation management measures, such as the definition of Marine Important Bird Areas for seabirds over the North Atlantic.     

Behavioral response of a generalist predator to chemotactile cues of two taxonomically distinct prey species

Chemotactile cues unintentionally left by animals can play a major role in predator–prey interactions. Specialized predators can use them to find their prey, while prey individuals can assess predation risk. However, little is known to date about the importance of chemotactile cues for generalist predators such as ants. Here, we investigated the response of a generalized predatory ant, Formica polyctena, to cues of two taxonomically distinct prey: a spider (Pisaura mirabilis) and a cricket (Nemobius sylvestris). In analogy, we studied whether crickets and spiders showed antipredator behavior in response to ant cues. When confronted with cues of the two prey species, Formica polyctena workers showed increased residence time and reduced movement speed, which suggests success-motivated searching behavior and thus increased foraging effort. The ants’ response did not differ between cues of the two prey species, coinciding with similar aggression and consumption rates of dead prey. However, the cuticular hydrocarbons, which likely resemble part of the potential cues, differed strongly between the species, with only few methyl-branched alkanes in common. This suggests that ants respond to multiple compounds left by other organisms with prey-search behavior. The two prey species, in turn, showed no detectable antipredator behavior in response to ant cues. Our study shows that ants can detect and respond to chemotactile cues of taxonomically and ecologically distinct prey species, probably to raise their foraging success. Using such chemotactile cues for prey detection may drastically increase their foraging efficiency and thus contribute to the high ecological success of ants.     

Heritability of the algal-foraging ability: an indirect benefit of female mate preference for males' carotenoid-based coloration in the guppy, Poecilia reticulata

Several sexual selection theories assume certain benefits of female mate preference. The direct benefit, i.e., the direct contribution from males to their offspring and females, has been well tested empirically. However, the indirect benefit, i.e., the male's genetic contribution to their offspring, has been poorly demonstrated. Female preference for males' carotenoid-based coloration is known in some animals. Since animals must acquire carotenoids through foods, it is often hypothesized that the brightness of the carotenoid-based coloration is a reliable indicator of the male's foraging ability. Hence, females' indirect benefits, such as greater foraging ability in their offspring, through mate preference for the carotenoid-based coloration are assumed. However, the heritability of the foraging ability for foods that serve as carotenoid resources has not been tested. In this study, a maze experiment was performed in guppies (Poecilia reticulata) to examine the heritability of the foraging ability for algae, carotenoid resources in nature. The latency for completing algal-foraging tasks in this experiment showed high individual variation. Heritable estimates of the foraging ability were substantial (h ² = 0.57 – 0.66) and significant, suggesting a genetic contribution to the foraging ability from fathers to their offspring. This result may support the hypothesis that indirect benefits influence the evolution of female choice.     

Path selection and foraging efficiency in Argentine ant transport networks

We experimentally investigated both individual and collective behavior of the Argentine ant Linepithema humile as they crossed symmetrical and asymmetrical bifurcations in gallery networks. Ants preferentially followed the branch that deviated the least from their current direction and their probability to perform a U-turn after a bifurcation increased with the turning angle at the bifurcation. At the collective level, colonies were better able to find the shortest path that linked the nest to a food source in a polarized network where bifurcations were symmetrical from one direction and asymmetrical from the other than in a network where all bifurcations were symmetrical. We constructed a model of individual behavior and showed that an individual’s preference for the least deviating path will be amplified via the ants’ mass recruitment mechanism thus explaining the difference found between polarized and non-polarized networks. The foraging efficiency measured in the simulations was three times higher in polarized than in non-polarized networks after only 15 min. We conclude that measures of transport network efficiency must incorporate both the structural properties of the network and the behavior of the network users.     

The chemosensory ability of the predatory leech Whitmania laevis (Arhynchobdellida: Haemopidae) for prey searching

Although prey-detecting and searching abilities of predatory leeches of rhynchobdellid or the Erpobdelliformes of arhynchobdellid species have been studied in the past, hirudiniformes leeches are rarely mentioned. In this study, we investigated the chemosensory ability for prey-detecting and searching in Whitmania laevis, a hirudiniformes species that mainly preys on freshwater snails, and examined if such ability aided in their prey selection. Five sympatric snail species, i.e., apple snail Pomacea canaliculata, thiarid snail Thiara tuberculata, viviparid snail Sinotaia quadrata, ear pond snail Radix auricularia swinhoei and tadpole snail Physa acuta were used as prey. Our results showed that W. laevis has the chemosensory ability to detect the waterborne odors of snails. However, they follow the snails by their mucus trails, and not by the odor that the snails leave in the water. Of these five snail species, W. laevis only followed the trails of the thiarid snails, ear pond snails and tadpole snails, and did not show a different response to the trails produced by snails of different sizes. Our results suggest that W. laevis can use waterborne odors to detect the existence of prey. They rely on mucus trails to follow their preferred prey, but do not distinguish between snails of a preferred size by their mucus trails. In addition, when following the trail of a preferred snail, W. laevis exhibits a newly described searching behavior, i.e., head tapping, and may use it to locate a snail trail and increase its probability of finding the trail-laying snail nearby.     

The importance of incorporating functional habitats into conservation planning for highly mobile species in dynamic systems

The distribution of mobile species in dynamic systems can vary greatly over time and space. Estimating their population size and geographic range can be problematic and affect the accuracy of conservation assessments. Scarce data on mobile species and the resources they need can also limit the type of analytical approaches available to derive such estimates. We quantified change in availability and use of key ecological resources required for breeding for a critically endangered nomadic habitat specialist, the Swift Parrot (Lathamus discolor). We compared estimates of occupied habitat derived from dynamic presence‐background (i.e., presence‐only data) climatic models with estimates derived from dynamic occupancy models that included a direct measure of food availability. We then compared estimates that incorporate fine‐resolution spatial data on the availability of key ecological resources (i.e., functional habitats) with more common approaches that focus on broader climatic suitability or vegetation cover (due to the absence of fine‐resolution data). The occupancy models produced significantly (P < 0.001) smaller (up to an order of magnitude) and more spatially discrete estimates of the total occupied area than climate‐based models. The spatial location and extent of the total area occupied with the occupancy models was highly variable between years (131 and 1498 km²). Estimates accounting for the area of functional habitats were significantly smaller (2–58% [SD 16]) than estimates based only on the total area occupied. An increase or decrease in the area of one functional habitat (foraging or nesting) did not necessarily correspond to an increase or decrease in the other. Thus, an increase in the extent of occupied area may not equate to improved habitat quality or function. We argue these patterns are typical for mobile resource specialists but often go unnoticed because of limited data over relevant spatial and temporal scales and lack of spatial data on the availability of key resources. Understanding changes in the relative availability of functional habitats is crucial to informing conservation planning and accurately assessing extinction risk for mobile resource specialists.     

Orientation and recruitment behavior in the ponerine ant Pachycondyla tesserinoda (Emery): laying of individual-specific trails during tandem running

Summary Tandem leaders of Pachycondyla tesserinoda mark their way during nest moving. While scouting and foraging for food in an unknown area, chemical orientation is used as well. The origin of the trail substance could not be identified. Secretions of gastral exocrine glands did not induce trail following behavior. Nor do these secretions elicit tandem-following reactions; rather the latter is released by the general body surface odor. The trail substance and the substance used for marking the nest entrance are colony-specific. Moreover, individual tandem leaders recognize and show a preference for their own trails. This extraordinary effect is independent of the age of the trail. P. tesserinoda workers search individually for food and new nest sites. Targets which are important for the colony are directly shown to nestmates by tandem running. Due to this type of foraging and scouting individual-specific trails may be advantageous for this ant.     

Behavioral responses to a novel predator and competitor of invasive mosquitofish and their non-invasive relatives (Gambusia sp.)

Attributes of the recipient community may affect the invasion success of arriving non-indigenous organisms. In particular, biotic interactions may enhance the resistance of communities to invasion. Invading organisms typically encounter a novel suite of competitors and predators, and thus their invasiveness may be affected by how they cope with these interactions. Behavioral plasticity may help invaders to respond appropriately to novelty. We examined the behavioral responses of highly invasive mosquitofish to representative novel competitors and predators they might encounter as they spread through North America. We compared the behavior of invasive Gambusia holbrooki and G. affinis to that of two close relatives of lower invasive potential (G. geiseri and G. hispaniolae) in order to elucidate whether responses to novelty related to invasiveness. In short-term assays, female Gambusia were paired with a novel competitor, Pimephales promelas, and a novel predator, Micropterus dolomieu. Behavioral responses were measured in terms of foraging success and efficiency, activity, refuge use, predator inspections, and interspecific aggression. Contrary to a priori predictions, invasive and non-invasive responses to novel interactions did not differ consistently. In response to novel competition, both invasive species increased foraging efficiency, but so did G. geiseri. In response to novel predation, only G. holbrooki decreased consumption and activity and increased refuge use. No antipredator response was observed in G. affinis. We found consistent differences, however, between invasives and non-invasives in foraging behavior. Both in the presence and absence of the competitor and the predator, invasives foraged more efficiently and consumed more prey than non-invasives.Communicated by P. Bednekoff     

The effect of hunger on the strength and duration of the antipredator behavioral response of green frog (<Emphasis Type="Italic">Rana clamitans</Emphasis>) tadpoles

The activity level of prey reflects a trade-off between predation risk and foraging gain. A number of theoretical and empirical studies have shown that a prey's energetic state or the level of its resource should influence this trade-off (i.e., what the optimal activity level at a level of predation risk is). Here, I show that the energetic state of prey may also influence the duration of their antipredator behavioral response. Green frog tadpoles (Rana clamitans) reduced their activity level for a shorter time during exposure to the chemical cue of predatory larval dragonflies (Anax spp.) as their time since last feeding increased (i.e., as their energetic state decreased). Interestingly, the tadpoles strongly reduced their activity level upon cue exposure in all treatments. Thus, the relative activity level of tadpoles at different energetic states varied over time.     

Vehicle pollution toxicity induced changes in physiology,defence system and biochemical characteristics of Calotropis procera L.

The present study aims to examine the physiological, biochemical and defence system responses of Calotropis procera to vehicle exhaust pollution. We selected various sample sites along two major roads in the Punjab province of Pakistan, i.e. Faisalabad to Sargodha road (FSR) and Pindi Bhattian to Lillah motorway (M-2). Traffic density at all sites and plant responses (i.e. chlorophyll a, chlorophyll b, total chlorophyll and carotenoid contents, transpiration rate, stomatal conductance, photosynthetic rate, sub-stomatal CO₂ concentration, water use efficiency, total free amino acids, total soluble proteins and total antioxidant activity) were measured. Levels of carbon (C) and nitrogen (N) and four metals of most possible concern – i.e. lead (Pb), zinc (Zn), cadmium (Cd) and nickel (Ni) – were also examined in all samples. We found their considerable deposition in all the samples along roads. Inhibitory effects of roadside pollutants were noted for photosynthetic pigments, stomatal conductance, transpiration rate, photosynthetic rate and total soluble proteins, whereas stimulatory effects were noted for sub-stomatal CO₂ concentration, free amino acids and total antioxidant activity. The stimulation of antioxidant enzymes activity revealed stress and mitigation of reactive oxygen species. The present study clearly signifies that C. procera has great potential to endure the stress caused by roadside pollutants.     

Costs of trail construction and maintenance in the leaf-cutting ant Atta columbica

Leaf-cutting ants of the genus Atta use trunk trails during foraging which may persist for months or years. The time and energy costs of trail construction and maintenance were estimated for colonies of Atta columbica on Barro Colorado Island, Panama, to determine if these costs are likely to constrain new trail construction and promote persistence of existing trails. Large workers 2.2-2.9 mm in headwidth participated in trail-clearing significantly more frequently than typical leaf-carriers, indicating that they may form a distinctive task group within the foraging force. Small litter items were carried off trails, while large ones were cut up before removal, greatly increasing the costs of removing large litter items. The average time cost of removing a kilogram of litter was estimated at 3,359 ant-hours, and energy costs at 4.6 kJ. Colonies maintained trail systems 267 m in length and 16.5 m² in area, and built an estimated 2.7 km of trail with an area of 134 m² during a year. Based on litter standing crop and estimates of litterfall rates, total costs to colonies averaged 11,000 ant-days of work and the energy equivalent of 8,000 leaf burdens. These costs are small relative to the number of available workers and rates of mass harvest, suggesting that costs do not significantly constrain trail construction. Instead, trails may persist because they provide access to high-quality resources or because only a few trails are required to fully exploit the foraging territory.     

The social network structure of a wild meerkat population: 2. Intragroup interactions

Knowledge of the structure of networks of social interactions is important for understanding the evolution of cooperation, transmission of disease, and patterns of social learning, yet little is known of how environmental, ecological, or behavioural factors relate to such structures within groups. We observed grooming, dominance, and foraging competition interactions in eight groups of wild meerkats (Suricata suricatta) and constructed interaction networks for each behaviour. We investigated relationships between networks for different social interactions and explored how group attributes (size and sex ratio), individual attributes (tenure of dominants), and ecological factors (ectoparasite load) are related to variation in network structure. Network structures varied within a group according to interaction type. Further, network structure varied predictably with group attributes, individual attributes, and ecological factors. Networks became less dense as group size increased suggesting that individuals were limited in their number of partners. Groups with more established dominant females were more egalitarian in their grooming and foraging competition interactions, but more despotic in their dominance interactions. The distribution of individuals receiving grooming became more skewed at higher parasite loads, but more equitable at low parasite loads. We conclude that the pattern of interactions between members of meerkat groups is not consistent between groups but instead depends on general attributes of the group, the influence of specific individuals within the group, and ecological factors acting on group members. We suggest that the variation observed in interaction patterns between members of meerkat groups may have fitness consequences both for individual group members and the group itself.     

Group size and foraging efficiency in yellow baboons

Summary I studied the foraging behaviour of adults in three different-sized groups of yellow baboons (Papio cynocephalus) at Amboseli National Park in Kenya to assess the relationship between group size and foraging efficiency in this species. Study groups ranged in size from 8 to 44 members; within each group, I collected feeding data for the dominant adult male, the highest ranking pregnant female, and the highest ranking female with a young infant. There were no significant differences between groups during the study in either the mean estimated energy value of the food ingested per day for each individual (385±27 kJ kg^-1 day^-1) or in the estimated energy expended to obtain that food (114±3 kJ kg^-1 day^-1). Mean foraging efficiency ratios, which reflect net energy gain per unit of foraging time, also did not vary as a function of the size of the group in which the baboons were living. There was substantial variation between days in the efficiency ratios of all animals; this was the result of large differences in energy intake rather than in the energy expended during foraging itself. The members of the smallest group spent on the average only one-half as much time feeding each day as did individuals in the two larger groups. However, they obtained almost as much energy while foraging, primarily because their rate of food intake while actually eating tended to be higher than the rate in the other groups. The baboons in the small group were observed closer to trees that they could climb to escape ground predators, and they also were more likely to sit in locations elevated above the ground while resting. Such differences would be expected if the members of the small group were less able to detect approaching predators than individuals that lived in the larger groups. The results of this study suggest that predator detection or avoidance, rather than increased foraging efficiency, may be the primary benefit of living in larger groups in this population.     

The interplay between foraging mode, habitat structure, and predator presence in antlions

Antlion larvae are sand-dwelling insect predators, which ambush small arthropod prey while buried in the sand. In some species, the larvae construct conical pits and are considered as sit-and-wait predators which seldom relocate while in other species, they ambush prey without a pit but change their ambush site much more frequently (i.e., sit-and-pursue predators). The ability of antlion larvae to evade some of their predators which hunt them on the sand surface is strongly constrained by the degree of sand stabilization or by sand depth. We studied the effect of predator presence, predator type (active predatory beetle vs. sit-and-pursue wolf spider), and sand depth (shallow vs. deep sand) on the behavioral response of the pit building Myrmeleon hyalinus larvae and the sit-and-pursue Lopezus fedtschenkoi larvae. Predator presence had a negative effect on both antlion species activity. The sit-and-wait M. hyalinus larvae showed reduced pit-building activity, whereas the sit-and-pursue L. fedtschenkoi larvae decreased relocation activity. The proportion of relocating M. hyalinus was negatively affected by sand depth, whereas L. fedtschenkoi was negatively affected also by the predator type. Specifically, the proportion of individual L. fedtschenkoi that relocated in deeper sand was lower when facing the active predator rather than the sit-and-pursue predator. The proportion of M. hyalinus which constructed pits decreased in the presence of a predator, but this pattern was stronger when exposed to the active predator. We suggest that these differences between the two antlion species are strongly linked to their distinct foraging modes and to the foraging mode of their predators. Reut Loria and Inon Scharf contributed equally to the paper.     

Energetics,reproductive suppression and obligate communal breeding in carnivores

Summary Leptogenys processionalis Jerdon forages on termites and other arthropods by raiding in branched trails. Growth and topology of these search trails were studied using Horton's (1945) technique orginally developed to analyze the branching pattern of river systems. Branching was always a bifurcating process and branches emerged symmetrically on either side of the main trail. Branching coefficients (R _b ) were similar to those of a few biological branching systems, such as lungs, that are considered to be non-random in their branching pattern. The R _b values indicated that the rate of branching and growth of trails remained constant within each foraging bout. The length of trails became shorter as they grew out and branched. The branching process was a function of the spatial separation of food patches in the terminal search field. Ants in the terminal search field send signals on encountering prey. The recruits cannot discriminate between these signals if they arise from two food patches situated <40 cm from each other, and hence converge on them in a single trail. However, discrimination is possible when food patches are >40 cm apart and hence recruits congregate on them separately in two trails, resulting in branching. Thus, the branching process is a result of independent decision by the ants conforming to certain simple rules and not a collective decision of the whole colony. We argue that mass recruiting ants selected to forage by branching pattern of trails because of its efficiency over other topologies (Stevens 1973) in minimizing the cost of travel, both from the nest to the food patches and between food patches. Further, the branch angles appear to be a trade-off to minimize travel cost and the resistance to the flow of ants comprising the column.Offprint requests to: T. Veena