Vibrational communication between hitchhikers and foragers in leaf-cutting ants (Atta cephalotes)

Abstract In a foraging column of the leaf-cutting ant Atta cephalotes, minim workers (the smallest worker subcaste) hitchhike on leaf fragments carried by larger workers. It has been demonstrated that they defend leaf carriers against parasitic phorid flies. The present study examines the cues used by the potential hitchhikers to locate leaf carriers. As recently reported, foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations serve as closerange recruitment signals. We tested the hypothesis that these plant-borne stridulatory vibrations are used by the potential hitchhikers to locate workers engaged in cutting. Three different lines of evidence support this view. Firstly, the repetition rate of the stridulations produced by foraging workers increases significantly as foragers maneuver the leaf fragment into the carrying position and walk loaded to the nest. This is the moment when hitchhikers usually climb on the leaf. Although the leaf-borne stridulatory vibrations are considerably attenuated when transmitted through the workers' legs, they can nevertheless be detected at short distances by minims. This subcaste is several times more sensitive to substrate-borne vibrations than larger workers. Secondly, when a stridulating and a silent leaf were simultaneously presented at the foraging site, minim workers spent significantly more time on the stridulating than on the silent leaf. Thirdly, hitchhiking was more frequent in leaf carriers which cut fragments out of the stridulating leaf than in those cutting the silent leaf.Abstract In a foraging column of the leaf-cutting ant Atta cephalotes, minim workers (the smallest worker subcaste) hitchhike on leaf fragments carried by larger workers. It has been demonstrated that they defend leaf carriers against parasitic phorid flies. The present study examines the cues used by the potential hitchhikers to locate leaf carriers. As recently reported, foraging workers stridulate while cutting a leaf fragment, and the stridulatory vibrations serve as closerange recruitment signals. We tested the hypothesis that these plant-borne stridulatory vibrations are used by the potential hitchhikers to locate workers engaged in cutting. Three different lines of evidence support this view. Firstly, the repetition rate of the stridulations produced by foraging workers increases significantly as foragers maneuver the leaf fragment into the carrying position and walk loaded to the nest. This is the moment when hitchhikers usually climb on the leaf. Although the leaf-borne stridulatory vibrations are considerably attenuated when transmitted through the workers' legs, they can nevertheless be detected at short distances by minims. This subcaste is several times more sensitive to substrate-borne vibrations than larger workers. Secondly, when a stridulating and a silent leaf were simultaneously presented at the foraging site, minim workers spent significantly more time on the stridulating than on the silent leaf. Thirdly, hitchhiking was more frequent in leaf carriers which cut fragments out of the stridulating leaf than in those cutting the silent leaf.Communicated by P. Pamilo     

Energetic and time costs of foraging in harvester ants,Pogonomyrmex occidentalis

Summary Western harvester ants, Pogonomyrmex occidentalis, preferentially utilize low vegetational cover pathways. Energetic costs for foraging ants were less than 0.1% of caloric rewards of harvested seeds, suggesting that reduction of energetic cost is not a major benefit of this preference. Walking speed was significantly faster on lower cover routes, increasing net return rates from equidistant artificial food sources. Undisturbed foragers on low cover routes traveled farther, increasing their total foraging area without increasing foraging time. These results suggest that in animals with low costs of locomotion relative to energetic rewards, time costs are more important than direct energetic costs in influencing foraging decisions. In baited experiments with equidistant food sources, preferential use of low cover routes resulted in a large increase in net energetic gain rate, but only a slight increase in energetic efficiency. Under natural conditions, net energetic gain rates were constant for foragers using low and high vegetational cover routes, but foragers using low cover paths had lower efficiencies. This suggests that net energetic gain rate is a more important currency than energetic efficiency for foraging harvester ants.     

Black yeast symbionts compromise the efficiency of antibiotic defenses in fungus-growing ants

Multiplayer symbioses are common in nature, but our understanding of the ecological dynamics occurring in complex symbioses is limited. The tripartite mutualism between fungus-growing ants, their fungal cultivars, and antibiotic-producing bacteria exemplifies symbiotic complexity. Here we reveal how black yeasts, newly described symbionts of the ant-microbe system, compromise the efficiency of bacteria-derived antibiotic defense in fungus-growing ants. We found that symbiotic black yeasts acquire nutrients from the ants' bacterial mutualist, and suppress bacterial growth. Experimental manipulation of ant colonies and their symbionts shows that ants infected with black yeasts are significantly less effective at defending their fungus garden from Escovopsis, a prevalent and specialized pathogen. The reduction of mutualistic bacterial biomass on ants, likely caused by black yeast symbionts, apparently reduces the quantity of antibiotics available to inhibit the garden pathogen. Success of the ant-fungal mutualism is directly dependent on fungus garden health. Thus our finding that black yeasts compromise the ants' ability to deal with the garden parasite indicates that it is an integral component of the symbiosis. This is further evidence that a full understanding of symbiotic associations requires examining the direct and indirect interactions of symbionts in their ecological community context.     

Use of stridulation in foraging leaf-cutting ants: mechanical support during cutting or short-range recruitment signal?

Foraging leaf-cutting ant workers stridulate while cutting a leaf fragment. Two effects of stridulation have recently been identified: (i) attraction of nestmates to the cutting site, employing substrate-borne stridulatory vibrations as short-range recruitment signals, and (ii) mechanical facilitation of the cut via a vibratome-effect. We asked whether foragers actually stridulate to support their cutting behavior, or whether the mechanical facilitation is an epiphenomenon correlated with the use of stridulation as recruitment signal. To differentiate between the two alternatives, workers of two different Atta species were presented with tender leaves of invariant physical traits, and their motivation to initiate recruitment was manipulated by varying the palatability of the leaves and the starvation of the colony. The lower the palatability of the harvested leaves, the lower the percentage of workers that stridulated while cutting, irrespective of the leaf’s physical features. After intense feeding, no workers were observed to stridulate while cutting tender leaves, and the percentage of stridulating workers increased with deprivation time. The results support the hypothesis that leaf-cutting ant workers stridulate during cutting in order to recruit nestmates, and that the observed mechanical facilitation of stridulation is an epiphenomenon of recruitment communication. Received: 25 January 1996/Accepted after revision: 13 July 1996     

The origin of the chemical profiles of fungal symbionts and their significance for nestmate recognition in <Emphasis Type="Italic">Acromyrmex</Emphasis> leaf-cutting ants

Cuticular hydrocarbon profiles are essential for nestmate recognition in insect societies, and quantitative variation in these recognition cues is both environmentally and genetically determined. Environmental cues are normally derived from food or nest material, but an exceptional situation may exist in the fungus-growing ants where the symbiotic fungus garden may be an independent source of recognition compounds. To investigate this hypothesis, we quantified the chemical profiles of the fungal symbionts of 18 sympatric colonies of Acromyrmex echinatior and Acromyrmex octospinosus and evaluated the quantitative variation of the 47 compounds in a multivariate analysis. Colony-specific chemical profiles of fungal symbionts were highly distinct and significantly different between the two ant species. We also estimated the relative genetic distances between the fungal symbionts using amplified fragment length polymorphism (AFLP) and correlated these with the overall (Mahalanobis) chemical distances between the colony-specific profiles. Despite the standardized laboratory conditions, the correlations were generally weak, but a statistically significant portion of the total variation in chemical profiles could be explained by genetic differences between the fungal symbionts. However, there was no significant effect of ant species in partial analyses because genetic differences between symbionts tend to coincide with being reared by different ant species. However, compound groups differed significantly with amides, aldehydes, and methyl esters contributing to the correlations, but acetates, alkanes, and formates being unrelated to genetic variation among symbionts. We show experimentally that workers that are previously exposed to and fed with the fungal symbiont of another colony are met with less aggression when they are later introduced into that colony. It appears, therefore, that fungus gardens are an independent and significant source of chemical compounds, potentially contributing a richer and more abundant blend of recognition cues to the colony “gestalt” than the innate chemical profile of the ants alone. Freddie-Jeanne Richard and Michael Poulsen contributed equally to this work.     

Anal gland secretion codes for family membership in the beaver

We investigated how information about family membership is coded by the individually specific anal gland secretion (AGS) in the beaver, Castor canadensis. Because beavers live in strict family units and relatives share more features in the AGS profile than non-relatives, family members share more AGS features than non-family members. Therefore, family recognition seems to be a natural consequence of the more specific kin recognition. Multivariate statistical analyses revealed two possibilities in coding for family membership using the multi-component AGS: either two (female) to three (male) AGS compounds or many compounds were used, but we were not able to determine which possibility is more likely. Compounds were not equally important in coding family membership information, and the interactions among compounds were complex. We attempted to reconstruct a lineage tree for the relationship among different families using two–three or many compounds. We found there was no significant difference for trees constructed by using few or many compounds in either males or females. However, the trees derived from male compounds and female compounds lacked good congruence. Received: 10 December 1998 / Accepted after revision: 13 July 1998     

The multifunctional pygidial gland secretion of the Steninae (Coleoptera: staphylinidae): ecological significance and evolution

The pygidial gland secretion of the rove beetle genera Stenus Latreille and Dianous Leach is composed of pyridine and piperidine-derived alkaloids and several terpene compounds. Two-choice bioassays with ants and fish, as well as agar diffusion assays, revealed that the secretion compounds are capable of deterring predators and protecting the beetles from infestation with microorganisms. In addition, the beetles use the secretion for rapid movements on the water surface, a process called skimming. Thus, originally developed to chemically defend the sensitive unprotected abdomen from predator attacks, the secretion of recent Steninae can be designated as multifunctional. Four of the alkaloid compounds occur as different configurational isomers in the secretion. Two-choice tests showed that ants discriminate between stereoisomers of stenusine, while there was no effect visible on bacteria in agar diffusion assays. Furthermore, there are evolutionary trends within the Steninae concerning the secretion composition, as some of the alkaloids primarily occur in phylogenetically basal species, while others are mainly restricted to derived species.     

Both evaluation of resource quality and speed of recruited leaf-cutting ants (Acromyrmex lundi) depend on their motivational state

Summary During recruitment, running velocity of both outbound and laden workers of the leaf-cutting ant Acromyrmex lundi depended on the information about resource quality they received from the first successful recruiter. In independent assays, single scout ants were allowed to collect sugar solutions of different concentrations and to recruit nestmates. Recruited workers were presented with standardized paper discs rather than the sugar solution given to the original recruiting ant. Outbound recruited workers were observed to run faster the more concentrated the solution found by the recruiter. Speed of disc-laden workers also depended on the concentration of the solution found by the recruiter, i.e. on the information about food quality they received, since they had no actual contact with the sugar solution. Disc-laden workers ran, as intuitively expected, slower than outbound workers. The reduction in speed, however, could not be attributed to the effects of the load itself, because workers collecting discs of the same weight, but with added sugar, ran as rapidly as outbound, unladen workers. Workers collecting standardized sugared discs reinforced the chemical trail on their way to the nest. The percentage of trail-layers was higher when workers were recruited to 10% than to 1% sugar solution, even though they collected the same kind of discs at the source. Their evaluation of resource quality, therefore, depended on their motivational state, which was modulated by the information they received during recruitment. Using previously published data on energetics of locomotion in leaf-cutting ants, travel costs of A. lundi workers recruited to sugar solutions of different concentration could be estimated. For workers recruited to the more concentrated solution, both speed and oxygen consumption rate increased by a roughly similar factor. Therefore, although workers ran faster to the high-quality resource, their actual energy investment per trip remained similar to that made by workers recruited to the low-quality resource. It is suggested that the more motivated workers reduced travel time without increasing energy costs during the trip. The adaptive value of these responses seems to be related to a rapid transmission of information about a newly discovered food source.     

Behavioural evidence for the involvement of Dufour’s gland secretion in nestmate recognition in the social wasp Polistes dominulus (Hymenoptera: Vespidae)

The results of recent studies by our group have suggested that in social wasps of the genus Polistes the Dufour’s gland is involved in kin recognition. In fact, the same hydrocarbons occurring on the cuticle are found in the gland secretion, and in P. dominulus the composition of the glandular secretion is more similar in foundresses belonging to the same colonies than in heterocolonial foundresses. In this study, P. dominulus colonies were experimentally presented with previously treated dead conspecific females. These lures had been deprived of their epicuticular lipids and coated with epicuticular or Dufour’s gland secretion extracts from females hetero- or homocolonial with respect to the tested colonies. The behaviour of the colonies towards these lures indicates that, like the epicuticular lipids, the Dufour’s gland secretion is involved in nestmate recognition. Received: 14 August 1995/Accepted after revision: 5 November 1995     

Experimental evidence for herbivore limitation of the treeline

The treeline ecotone divides forest from open alpine or arctic vegetation states. Treelines are generally perceived to be temperature limited. The role of herbivores in limiting the treeline is more controversial, as experimental evidence from relevant large scales is lacking. Here we quantify the impact of different experimentally controlled herbivore densities on the recruitment and survival of birch Betula pubescens tortuosa along an altitudinal gradient in the mountains of southern Norway. After eight years of summer grazing in large-scale enclosures at densities of 0, 25, and 80 sheep/km2, birch recruited within the whole altitudinal range of ungrazed enclosures, but recruitment was rarer in enclosures with low-density sheep and was largely limited to within the treeline in enclosures with high-density sheep. In contrast, the distribution of saplings (birch older than the experiment) did not differ between grazing treatments, suggesting that grazing sheep primarily limit the establishment of new tree recruits rather than decrease the survival of existing individuals. This study provides direct experimental evidence that herbivores can limit the treeline below its potential at the landscape scale and even at low herbivore densities in this climatic zone. Land use changes should thus be considered in addition to climatic changes as potential drivers of ecotone shifts.     

Allometry and the geometry of leaf-cutting in Atta cephalotes

Summary This study considers the relationship of both leg length and the geometry of leaf-cutting to load-size determination by the highly polymorphic leaf-cutting ant Atta cephalotes. A. cephalotes workers anchor on the leaf edge by their hind legs and pivot around them while cutting arcs from leaves. I tested the hypothesis that, for an ant cutting a semicircular leaf fragment, fragment area is determined by a fixed reach while cutting. This reach hypothesis predicts that ants should cut the same fragment-area for at all leaf types. Also, if the radius of the semicircular fragment is proportional to hind leg length, this hypothesis predicts that leaf area should be proportional to hind-leg length squared. The field work was carried out in March–April 1990 and June 1991 in Heredia Province, Costa Rica. I measured hind-leg length for workers of different masses. I then measured leaf-fragment area and mass for workers cutting semicircular fragments from leaves of different densities (mass/area). The logarithmic relationship between ant mass (M _a) and hind-leg length L accelerated negatively (Fig. 1). As a result of this complex allometry, relative leg length (L/M _a ^0.33) increased with ant mass up to a mass of 7.4 mg. Above 7.4 mg, relative leg length decreased. For foragers cutting semicircular fragments, the area cut by an ant of a given size showed no significant difference among leaves of different densities (Fig. 2). Leaf area (A) increased as a function of leg length to the 1.9 power (Fig. 4), an exponent not significantly different from the square function expected if the radius of a fragment is determined by the ant's reach. As a result of this consistent mode of fragment-area determination, the mass of fragments cut by an ant of a given size was significantly greater when cutting denser leaves (Fig. 3) and relative area (A/M _a) cut decreased with increasing ant mass. However, because larger ants generally cut denser leaves (Table 1), the increased density of thick leaves was offset by the reduced relative area cut by the larger ants. Overall, 93% of the foragers cut fragments weighing between 1.5 and 6 times their own body mass (Table 1). Earlier studies found that this broad load-mass range maximized the biomass-transport rate (mass/distance/time) and transport efficiency (mass/distance/energy cost). Thus, A. cephalotes does not solve the problem of matching ant mass and load mass at leaves of different densities with flexibility in the leaf-cutting behavior of individual ants. Instead, individual ants employ a single simple behavioral rule, but workers of different sizes and body proportions tend to cut leaves of different densities.     

Experimental evidence for internal predation in microzooplankton communities

The fate of microzooplankton production, whether it is channeled to mesozooplankton or recycled within the microbial food web, has major implications for the oceanic carbon cycle. The aim of this study was to estimate internal predation within naturally occurring microzooplankton communities. A dilution series based on the Landry and Hasset technique was created by mixing 200-μm-screened water (used as whole water) with 5-μm-screened seawater due to the dominance of pico- and small nanoplankton at our study site. This modification of the original technique allows for gradual reduction in microzooplankton abundance and thus internal predation while maintaining sufficient phytoplankton prey levels for microzooplankton growth in diluted treatments. Microzooplankton growth and mortality rates were calculated based on the changes in abundance during 24-h incubation. In the diluted treatments, microzooplankton growth rates were significantly higher (1.21 ± 0.20 day^?1 for ciliates and 0.88 ± 0.05 day^?1 for heterotrophic dinoflagellates) compared to those in whole seawater where microzooplankton abundance remained unchanged or even declined over time. Approximately 79 % of microzooplankton production was consumed within the microzooplankton, with aloricate ciliates being the most vulnerable to predation. These findings support the assumption that trophic interactions between microzooplankton can be an important factor controlling their production and, thus, energy transfer in picoplankton-dominated pelagic ecosystems.     

The costs and benefits of cooperation between the Australian lycaenid butterfly,Jalmenus evagoras,and its attendant ants

Summary The larvae and pupae of the Australian lycaenid butterfly, Jalmenus evagoras associate mutualistically with ants in the genus Iridomyrmex. Four ant exclusion experiments in three field sites demonstrated that predation and parasitism of J. evagoras are so intense that individuals deprived of their attendant ants are unlikely to survive. Larvae and pupae of J. evagoras aggregate, and the mean number of attendant ants per individual increases with larval age and decreases with group size. Field observations showed that young larvae could gain more attendant ants per individual by joining the average size group of about 4 larvae than by foraging alone. Aggregation behaviour is influenced by ant attendance: young larvae and pupating fifth instars aggregated significantly more often on plants with ants than on plants where ants had been excluded. In return for tending and protecting the larvae, ants were rewarded by food secretions that can amount to as much as 409 mg dry biomass from a single host plant containing 62 larvae and pupae of J. evagoras over a 24 h period. Larval development in the laboratory lasted approximately a month, and larvae that were tended by ants developed almost 5 days faster than larvae that were not tended. However, tended individuals, particularly females, pupated at a significantly lower weight than their untended counterparts, and the adults that eclosed from these pupae were also lighter and smaller. On average, pupae that were tended by ants lost 25% more weight than untended pupae, and in contrast with larvae, they took longer to eclose than pupae that were not tended. These experimental results are discussed in terms of costs and benefits of association for both partners, and of aggregation for the lycaenids.     

Experimental evidence of reciprocal altruism in the pied flycatcher

Although human behaviour abounds with reciprocal altruism, few examples exist documenting reciprocal altruism in animals. Recent non-experimental evidence suggests that reciprocal altruism may be more common in nature than previously documented. Here we present experimental evidence of mobbing behaviour, the joint assault on a predator in an attempt to drive it away, as reciprocal altruism in the breeding pied flycatcher (Ficedula hypoleuca). Given a choice, pied flycatchers assisted in mobbing initiated by co-operating neighbours and did not join in mobbing when initiated by conspecific neighbours which had defected from necessary assistance 1 h before. The results suggest the birds followed a ‘tit-for-tat’-like strategy and that mobbing behaviour of breeding birds may be explained in terms of reciprocal altruism.     

Experimental evidence for mediation of competition by habitat succession

Habitat succession is thought to influence the importance of competition in assemblages. Competitive interactions are considered of critical importance in structuring ant assemblages, but field experiments show varied effects. I tested how succession in managed boreal forests affects the outcome of competition from dominant red wood ants, Formica aquilonia, through a removal experiment in replicated stands of three different ages (0-4, 30-40, and 80-100 years old). F. aquilonia abundance was reduced by 87%, and procedural controls showed no nontarget effects. The succession gradient revealed the full range of possible responses from ant species: decreases in 1-4-year-old stands, increases in 30-40-year-old stands, and no effects in 80-100-year-old stands, where diversity was lowest. Habitat succession thus regulates competitive interactions in this system. I propose a model for this system, where competitive effects depend on time since disturbance. In this case, soon after disturbance the dominant species facilitates increases in the abundance of other species. At intermediate times, competition reduces the abundance of some species. Finally, in long-undisturbed habitats, competitors may fail to evolve, particularly in high-stress environments. Interactions between competition and habitat succession may explain why structuring effects of ecologically dominant species appear inconsistent.     

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.     

Experimental evidence for direct nutrient assimilation by the lophophore of articulate brachiopods

The uptake of inorganic and non-nutritive organic molecules has been compared with uptake of nutritive molecules by the articulate brachiopod Terebratalia transversa (Sowerby). Only minimal uptake of Na₂ ¹⁴CO₃ and ¹⁴C-urea was observed, while ¹⁴C-glucose was concentrated extensively. After administration of a dilute solution of ¹⁴C-glucose over timed intervals, whole organ counts and autoradiographs showed that labelled material was accumulated along the exposed ciliated epidermal tissue of the lophophore and mantle and concentrated along the peritoneal lining of the coelom even before appearing in the gastrointestinal tract. The presence or absence of bacteria had little discernible effect upon extent and rate of uptake. The uptake experiments suggest that the lophophore not only creates an inhalent and exnalent current as is common in other filter-feeders, but also appears to be adapted for extraction of dilute nutrients in seawater. This ability of the lophophore to extract nutrients may help explain the evolutionary trend of the lengthening of the articulate lophophore and the reduction of the intestine to a short blind-ended gut.This work was supported by National Science Foundation Grant GB-20067.     

Experimental evidence for helping and hindering by previous offspring in the cooperative-breeding Seychelles warbler Acrocephalus sechellensis

Prebreeding Seychelles warblers (Acrocephalus sechellensis) frequently act as helpers on their natal territory, aiding in territory defence, predator mobbing, nestbuilding, incubation (only females) and feeding dependent young of their parents. In some cases helpers could attain breeding status (e.g. by joint-nesting) in their natal group and become co-breeders. Comparisons of group size and reproductive success on a given quality territory suggest that the presence of alloparents (helpers and cobreeders) significantly affects the reproductive success of their parents. The influence of alloparents on reproductive success was examined by removing alloparents from breeding units and comparing the success of natural-sized and artificially reduced groups. Removal experiments, controlled for territory quality, group size and breeder age, showed that the presence of one alloparent significantly improved the reproductive success of its parents. Analysis strongly suggests that this was entirely due to helping behaviour (i.e. providing care to offspring of their parents), thereby improving the helper's inclusive fitness benefits from staying at home. However, these experiments showed also that the presence of two or more alloparents in medium-quality territories significantly decreased reproductive success, compared with groups with one alloparent. Several lines of evidence suggest that this may have been due to the joint-nesting and reproductive competition that could occur in breeding groups, or simply to resource depression when a large number of previous offspring remained on their natal territory.     

An experimental study of competition between fire ants and Argentine ants in their native range

An understanding of why introduced species achieve ecological success in novel environments often requires information about the factors that limit the abundance of these taxa in their native ranges. Although numerous recent studies have evaluated the importance of natural enemies in this context, relatively few have examined how ecological success may result from differences in the magnitude of interference competition between communities in the native and introduced ranges of nonnative species. Here we examine how native-range competitive environments may relate to invasion success for two important invasive species, the red imported fire ant (Solenopsis invicta) and the Argentine ant (Linepithema humile), in a region of native-range sympatry. At two study sites in northern Argentina, we used stable-isotope analysis, a variety of observational approaches, and two different reciprocal removal experiments to test (1) whether S. invicta competes asymmetrically with L. humile (as suggested by the 20th century pattern of replacement in the southeastern United States) and (2) the extent to which these two species achieve behavioral and numerical dominance. Stable-isotope analysis and activity surveys indicated that S. invicta and L. humile are both omnivores and forage during broadly overlapping portions of the diel cycle. Short-term removal experiments at baits revealed no competitive asymmetry between S. invicta and L. humile. Longer-term colony removal experiments illustrated that S. invicta and L. humile experience an approximately equal competitive release upon removal of the other. Our results indicate that neither S. invicta nor L. humile achieves the same degree of behavioral or ecological dominance where they co-occur in native populations as they do in areas where either is common in their introduced range. These results strongly suggest that interspecific competition is an important limiting factor for both S. invicta and L. humile in South America.     

Foraging costs of a tail ornament: Experimental evidence from two populations of barn swallows Hirundo rustica with different degrees of sexual size dimorphism

Secondary sexual characters are assumed to be costly to produce or maintain. A test of this assumption was performed using the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject of a directional female mate preference. A possible cost of sexual signalling in male barn swallows arises from increased flight cost during foraging in this aerially insectivorous species. A longer tail may impose a greater drag during flight and thereby affect foraging ability. This was tested by determining the relationship between experimentally modified male tail lengths and number and size of prey delivered to offspring in Spain, where sexual size dimorphism in tail length is small, compared to Denmark, where dimorphism is large. Food boluses contained significantly fewer small insects in Spain than in Denmark. Males with elongated tails captured more and smaller insects while males with shortened tails captured fewer and larger prey items at both sites. Males with naturally long tails were less affected by experimental treatment in terms of effects on the number and the size of prey delivered to their offspring, a finding consistent with a long tail being a condition-dependent viability indicator. The effect of a given degree of tail manipulation on prey size and number of prey per bolus was larger in Spain than in Denmark. These results demonstrate that (1) tail length in male barn swallows affects foraging, and (2) larger sexual size dimorphism occurs where the foraging cost of an increment in ornament size is smallest.Communicated by M. Zuk