Individual inter‐annual nest‐site relocation behaviour drives dynamics of a recently established Barnacle Goose Branta leucopsis colony in sub‐arctic Russia

Avian breeding colonies are generally in a continuous state of flux, some parts growing whilst others shrink as individuals move within the colony on the search for better nest‐sites. We examined the role of experience in breeding patch choice by individually marked Barnacle Geese Branta leucopsis in a recently established colony in sub‐arctic Russia. Individuals failing to reproduce successfully tended to shift nest location further the following season than did successful pairs, and they did so towards the most dense nest aggregations within the colony, where reproductive success was higher. We suggest that individual decisions on nest‐site choice shape the spatial dynamics of this colony.     

Restoration and management for cliff‐nesting birds in Mediterranean mining sites: the Sand Martin case study

Habitat enhancement for birds is frequently implemented during mine site restoration. Cliff‐nesting birds often colonize anthropogenic environments such as mining areas (aggregate sites and quarries for aggregate and cement production). Mining activity can compromise breeding success, causing cliff‐nesting birds to depend on the management and restoration of mining areas. The objective of our study is to assess the importance of mine site habitats for Sand Martin conservation and reconcile mining activity with breeding success in Mediterranean environments. We studied Sand Martin breeding habitat preferences in mining areas at three spatial scales. At the mining site scale, we compared 10 mining sites with Sand Martin burrows with 19 mining sites without burrows. At the colony scale (vertical structures with colonies), we evaluated the relationships between the number of breeding pairs, number of burrows, and colony characteristics within 30 distinct Sand Martin colonies. At the burrow scale, we compared the characteristics of the available vertical structure with the areas used by Sand Martins. At the mining site scale, Sand Martins preferred more surface of water bodies, shorter distances to flowing water, older sites, and mining sites which produce aggregates instead of cement. At the colony scale, Sand Martins preferred southwest orientations and stockpiles to vertical extraction faces. At the burrow scale, birds preferred the most vertical areas of the face. Our results support the need for effective habitat restoration and improved management for more effective Sand Martin conservation within mining areas. Simple interventions can enhance habitat quality and conservation of cliff‐nesting birds.     

Factors affecting population regulation of a colonial vulture

Two hypotheses have been proposed to link population regulation to density‐dependent changes in demographical parameters: the habitat heterogeneity hypothesis (HHH) states that, as population density rises, an increasing proportion of individuals are forced to occupy low‐quality territories, which provokes a decline in average per‐capita survival and/or productivity although some individuals show no decline in fecundity; and the individual adjustment hypothesis (IAH), which suggests that increased densities lead to reductions in survival and/or fecundity by enhancing agonistic interactions, which affect all individuals to a similar extent. However, density‐dependent effects can be affected by density‐independent factors (DIF), such as weather. We test the effects of density dependence on annual reproductive success in Griffon Vultures Gyps fulvus at four spatial scales, nest‐site, cliff, colony and metacolony, in northern Spain from 2008 to 2015. Our results showed most support for the HHH at all scales. At the colony and cliff scale, IAH and DIF had similar importance, whereas there was little evidence of IAH at the metacolony and the nest scale. The best protected eyries (caves, potholes and sheltered ledges) produced the most fledglings and were used preferentially, whereas low‐quality eyries (exposed ledges or open crevices) were used only when the number of breeders increased. The significant interaction between breeding failure and density found for the more exposed eyries suggests that at higher densities, breeding pairs are forced to use poorer nesting areas, and the negative effect of density at the cliff scale could be due to the combined effect of a higher proportion of pairs using low‐quality eyries and the negative effect of rainfall.     

Nest survival is influenced by parental behaviour and heterospecifics in a mixed‐species colony

Studies of avian nest success often focus on examining influences of variation in environmental and seasonal factors. However, in‐depth evaluations can also incorporate variation in individual incubation behaviour to further advance our understanding of avian reproductive ecology. We examined these relationships in colonially nesting Black‐crowned Night‐Herons Nycticorax nycticorax using intensive video‐monitoring methods to quantify incubation behaviours. We modelled nest survival as a function of both extrinsic factors and incubation behaviours over a 3‐year period (2010–12) on Alcatraz Island, USA. Model‐averaged parameter estimates indicated that nest survival increased as a function of greater incubation constancy (% of time spent incubating eggs within a 24‐h period), and average daily precipitation throughout the nesting stage. Common Ravens Corvus corax are the only known nest predator of Night‐Herons on Alcatraz Island, as on many other coastal Pacific islands. We also investigated the effects of heterospecific nesting of California Gulls Larus californicus and Western Gulls Larus occidentalis in a mixed‐species colony with Night‐Herons, based on nesting proximity data collected over a 2‐year period (2011–12). This second analysis indicated that, in addition to incubation behaviours, nesting heterospecifics are an important factor for explaining variation in Night‐Heron nest survival. However, contrary to our original expectation, we found that Night‐Herons experienced increased nest survival with increasing distance from gull colony boundaries. These results may apply to other areas with multiple colonial nesting species and similar predator communities and climatic patterns.     

Cues of Predation Risk Induce Instar‐ and Genotype‐Specific Changes in Pea Aphid Colony Spatial Structure

Deploying collective antipredator behaviors during periods of increased predation risk is a major determinant of individual fitness for most animal groups. Pea aphids, Acyrthosiphon pisum, which live in aggregations of genetically identical individuals produced via asexual reproduction warn nearby conspecifics of pending attack by secreting a volatile alarm pheromone. This alarm pheromone allows clone‐mates to evade predation by walking away or dropping off the host plant. Here, we test how a single alarm pheromone emission influences colony structure and defensive behavior in this species. Relative to control colonies, groups exposed to alarm pheromone exhibited pronounced escape behavior where many individuals relocated to adjacent leaves on the host plant. Alarm pheromone reception, however, also had subtle instar‐specific effects: The proportion of 1st instars feeding nearest the leaf petiole decreased as these individuals moved to adjacent leaves, while the proportion of 2nd–3rd instars feeding nearest the leaf petiole remained constant. Fourth instars also dispersed to neighboring leaves after pheromone exposure. Lastly, alarm pheromone reception caused maternal aphids to alter their preferred feeding sites in a genotype‐specific manner: Maternal aphids of the green genotype fed further from the petiole, while maternal aphids of the pink genotype fed closer to the petiole. Together, our results suggest that aphid colony responses to alarm pheromone constitute a diversity of nuanced instar‐ and genotype‐specific effects. These behavioral responses can dramatically change the spatial organization of colonies and their collective defensive behavior.     

Influence of density‐dependent competition on foraging and migratory behavior of a subtropical colonial seabird

Density‐dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central‐place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density‐dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican (Pelecanus occidentalis), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density‐dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density‐dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density‐dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density‐dependent competition on breeding success and population regulation remain uncertain in this system.     

X‐ray computed tomography reveals that intraspecific competition promotes soldier differentiation in a one‐piece nesting termite

Investment in soldier production in eusocial lineages involves a trade‐off between maintenance costs and defense benefits. Termites are eusocial insects that live in colonies organized into three castes: primary reproductives, soldiers, and workers or pseudergates. Neotermes chilensis (Blanchard) (Isoptera: Kalotermitidae) is a one‐piece nesting termite that nests and forages in a single piece of wood. Two scenarios may be of importance in a defense context of one‐piece nesting termites: during swarms, when colonies may be invaded by winged termites (alates) in search of a place to found a new colony, and when colonies of conspecifics are present within the same substrate. It was hypothesized that the ratio of soldiers to non‐soldiers would be higher at the onset of the swarming period and in substrates bearing more than one termite colony. A method based on X‐ray computed tomography (CT) was developed to study gallery connectivity in colonies of N. chilensis and caste composition within colonies. Computed tomography allowed the digital reconstruction of the galleries within the substrate, even when they belonged to different colonies, and was effective in distinguishing termites from substrate, and soldiers from reproductives and pseudergates. Using CT, the ratio of soldiers to non‐soldiers was shown to be highest in colonies within multicolonial scapes (i.e., neighboring colonies were present in the same substrate) during the swarming season, thus supporting our initial hypotheses. These results constitute a unique example of induced defenses arising from intraspecific interactions in termites.     

Suburban sprawl: environmental features affect colony social and spatial structure in the black carpenter ant,Camponotus pennsylvanicus

1. In social insects, the number of nests that a colony inhabits may have important consequences for colony genetic structure, the number of queens, sex allocation, foraging efficiency, and nestmate recognition. Within the ants, colonies may either occupy a single nest (monodomy) or may be organised into a complex network of nests and trails, a condition known as polydomy. 2. The current study is a large‐scale, long‐term, comprehensive field examination of various features of colony social and spatial structure in the facultatively polydomous black carpenter ant, Camponotus pennsylvanicus (DeGeer). The study examined the density, persistence, and the spatiotemporal distribution of colonies across a gradient of land disturbance associated with urban development. The temporal and spatial pattern of nest use was compared between fragmented landscapes where nesting sites were interspersed among human‐built structures (urban plots) and less disturbed landscapes with higher tree density (suburban plots). In addition, nesting site fidelity and changes in colony spatial structure were monitored over 7 years. 3. Long‐term monitoring and extensive sampling over a large spatial area allowed the first comprehensive insight into the spatiotemporal dynamics of colony and population structure in C. pennsylvanicus. A total of 1113 trees were inspected over 233 ha. Camponotus pennsylvanicus were active on 348 of the 1113 trees (31%) and these represented 182 distinct colonies. The colonisation rate remained relatively stable over 7 years suggesting that an equilibrium point had been reached. Relative to the suburban plots, tree density was 65% lower in the urban plots. The proportion of trees colonised by C. pennsylvanicus was significantly higher in the urban plots suggesting that intraspecific competition for nesting sites may be especially high in areas with lower tree density. Colony spatial structure also differed significantly between habitats and a higher incidence of monodomy was observed in the urban environment. The average number of trees per colony across all subplots was 1.95 (range 1–4) indicating that C. pennsylvanicus are weakly polydomous. 4. The composite picture that emerges for C. pennsylvanicus colonies in the urban habitat is a chain reaction of events: (i) the urban habitat has a lower tree density, (ii) lower tree density results in higher tree colonisation rate, (iii) higher tree colonisation rate results in simpler colony spatial structure (i.e. higher incidence of monodomy), and (iv) simpler colony spatial structure results in numerically smaller colonies. Long‐term monitoring of the spatiotemporal pattern of nest site use in selected colonies revealed a unique trend. While worker counts in selected colonies remained relatively stable throughout the course of the study, colony spatial structure changed considerably with 28% of colonies experiencing a change. Furthermore, the likelihood of detecting a change in colony spatial structure increased with the amount of time passing from the initial inspection. 5. In conclusion, tree density has a significant effect on a number of important colony features in C. pennsylvanicus. Besides tree density, other environmental features such as human‐built structures cause habitat fragmentation and may act as natural barriers to worker dispersal and/or foraging. Such barriers may ultimately affect the social and/or spatial structure at both the colony and the population level.     

Individual and Group Personalities Characterise Consensus Decision‐Making in an Ant

Non‐human animals can exhibit idiosyncratic behaviour across individuals in much in the same way as humans. Animals with specific personalities may have advantages in some environments, and this idiosyncrasy may thus be of considerable ecological and evolutionary importance. In group‐living organisms, personality can occur at the level of the group as well as that of the individual. However, at present, we have very little understanding of the possible benefits of group‐level personality, and how this is linked with individual personality. In this study, I examine the influence of individual and group personality during the process of colony migration in the Japanese ant, Myrmecina nipponica. These ants use a consensus decision process to decide among alternatives when searching for a new home. Individuals contribute to this process by scouting for new nest sites, recruiting nestmates by laying pheromone trails, and carrying brood to the new site, although whether these roles are consistent among individuals and how roles are distributed within and between colonies remain unclear. Individual contributions to the nest‐site selection process were quantified over five repeated relocations in five colonies. Results demonstrate that contributions to the relocation effort were highly skewed within the colonies and that individuals were consistent in their contributions over repeated relocation events. Furthermore, the distribution of effort differed between colonies, indicating that intercolony differences in composition of behavioural types resulted in colony‐level personality. While these differences did not lead to any detectable difference in relocation performance between colonies in the simple experimental arrangement used, colony personality could influence decision outcomes in more complex environments.     

Determinants of between‐year burrow re‐occupation in a colony of the European bee‐eater Merops apiaster

Re‐occupation of existing nesting burrows in the European bee‐eater Merops apiaster has only rarely – and if so mostly anecdotically – been documented in the literature record, although such behavior would substantially save time and energy. In this study, we quantify burrow re‐occupation in a German colony over a period of eleven years and identify ecological variables determining reuse probability. Of 179 recorded broods, 54% took place in a reused burrow and the overall probability that one of 75 individually recognized burrows would be reused in a given subsequent year was estimated as 26.4%. This indicates that between‐year burrow reuse is a common behavior in the study colony which contrasts with findings from studies in other colonies. Furthermore, burrow re‐occupation probability declined highly significantly with increasing age of the breeding wall. Statistical separation of within‐ and between‐burrow effects of the age of the breeding wall revealed that a decline in re‐occupation probability with individual burrow age was responsible for this and not a selective disappearance of burrows with high re‐occupation probability over time. Limited duty cycles of individual burrows may be caused by accumulating detritus or decreasing stability with increasing burrow age. Alternatively, burrow fidelity may presuppose pair fidelity which may also explain the observed restricted burrow reuse duty cycles. A consequent next step would be to extend our within‐colony approach to other colonies and compare the ecological circumstances under which bee‐eaters reuse breeding burrows.     

Polygynous supercolonies of the acacia‐ant Pseudomyrmex peperi,an inferior colony founder

In ant–plant protection mutualisms, plants provide nesting space and nutrition to defending ants. Several plant–ants are polygynous. Possessing more than one queen per colony can reduce nestmate relatedness and consequently the inclusive fitness of workers. Here, we investigated the colony structure of the obligate acacia‐ant Pseudomyrmex peperi, which competes for nesting space with several congeneric and sympatric species. Pseudomyrmex peperi had a lower colony founding success than its congeners and thus, appears to be competitively inferior during the early stages of colony development. Aggression assays showed that P. peperi establishes distinct, but highly polygynous supercolonies, which can inhabit large clusters of host trees. Analysing queens, workers, males and virgin queens from two supercolonies with eight polymorphic microsatellite markers revealed a maximum of three alleles per locus within a colony and, thus, high relatedness among nestmates. Colonies had probably been founded by one singly mated queen and supercolonies resulted from intranidal mating among colony‐derived males and daughter queens. This strategy allows colonies to grow by budding and to occupy individual plant clusters for time spans that are longer than an individual queen’s life. Ancestral states reconstruction indicated that polygyny represents the derived state within obligate acacia‐ants. We suggest that the extreme polygyny of Pseudomyrmex peperi, which is achieved by intranidal mating and thereby maintains high nestmate relatedness, might play an important role for species coexistence in a dynamic and competitive habitat.     

All you can eat: is food supply unlimited in a colonially breeding bird?

Food availability is generally considered to determine breeding site selection and therefore plays an important role in hypotheses explaining the evolution of colony formation. Hypotheses trying to explain why birds join a colony usually assume that food is not limited, whereas those explaining variation in colony size suggest that food is under constraint. In this study, we investigate the composition and amount of food items not eaten by the nestlings and found in nest burrows of colonially nesting European bee‐eaters (Merops apiaster). We aimed to determine whether this unconsumed food is an indicator of unlimited food supply, the result of mistakes during food transfer between parents and chicks or foraging selectivity of chicks. Therefore, we investigated the amount of dropped food for each nest in relation to reproductive performance and parameters reflecting parental quality. Our data suggest that parents carry more food to the nest than chicks can eat and, hence, food is not limited. This assumption is supported by the facts that there is a positive relationship between dropped food found in a nest and the number of fledglings, nestling age, and chick health condition and that the amount of dropped food is independent of colony size. There is variation in the amount of dropped food within colonies, suggesting that parent foraging efficiency may also be an important determinant. Pairs nesting in the center of a colony performed better than those nesting on the edge, which supports the assumption that quality differences between parents are important as well. However, dropped food cannot be used as an indicator of local food availability as (1) within‐colony variation in dropped food is larger than between colony variation and, (2) the average amount of dropped food is not related to colony size.     

Detectability and visibility biases associated with using a consumer‐grade unmanned aircraft to survey nesting colonial waterbirds

Surveys of colonial‐nesting waterbirds are needed to assess population trends and gain insight into the health of wetland ecosystems. Use of unmanned aerial systems (UAS) for such surveys has increased over the past decade, but possible sources of bias in surveys conducted with UAS have not been examined. We examined possible visibility biases associated with using a UAS to survey waterbird colonies in cypress‐tupelo watersheds and coastal island habitats in Texas in 2016. We used known numbers of four waterbird decoy types, including Black Skimmers (Rynchops niger), terns, and white‐ and dark‐plumaged herons, to estimate their detectability in each habitat. Six observers independently counted decoys from aerial imagery mosaics taken with a consumer‐grade, off‐the‐shelf quadcopter drone. We used generalized linear mixed‐effects models to estimate detection probabilities of each decoy type. Black Skimmers at the coastal island had a detection probability of just 53%. Detectability of both white‐ and dark‐plumaged herons was lower in the canopied cypress‐tupelo habitat than the coastal island. In addition, cloud cover > 50% further reduced detectability of white heron decoys in cypress‐tupelo habitat. Use of the double‐count method yielded biased‐low abundance estimates for white‐ and dark‐plumaged herons in canopied sites, suggesting that habitat differences were a greater source of bias than observer error. Black Skimmers were the only decoy type to be imperfectly detected at the coastal island, a surprising result given the stark contrast of their plumage with their sand and shell nesting substrate. Our results indicate that UAS‐derived photographic surveys are prone to low detection probabilities at sites where vegetation occludes nests. In habitats without canopy, however, UAS surveys show promise for obtaining accurate counts of terns, white herons, and dark herons.     

Spatial differences in breeding success in the pied avocet Recurvirostra avosetta: effects of habitat on hatching success and chick survival

I studied the breeding biology of pied avocets Recurvirostra avosetta in natural habitats (alkaline lakes), and in semi-natural sites (dry fishpond, reconstructed wetlands) in Hungary to relate habitat selection patterns to spatial and temporal variation in breeding success. Colonies were initiated earlier in semi-natural sites than in natural habitats, and earlier on islands than on the mainland. Hatching success was higher on islands than on the mainland, in semi-natural sites than in natural habitats, in colonies of at least 15 pairs than in smaller colonies, and for nests initiated earlier than later within a colony. Fledging success was higher in the wet years (1999–2000) than in the dry year (1998), decreased strongly by season in both habitats and increased with average daily temperature in the first week post-hatch in 1999–2000. Most pairs hatching young in semi-natural sites attempted to lead their chicks to feeding areas in natural habitats, whereas no such movement occurred in the opposite direction. Chick mortality due to predation was high during brood movements and only 23% of the pairs moving their young produced fledglings, compared to 43% for pairs remaining in semi-natural sites and 68% for pairs hatching and rearing young in natural habitats (total n=192 broods). These results suggest that semi-natural sites were more suitable for nesting, whereas natural habitats were more suitable for chick-rearing. The opposing trends in habitat-related breeding success between nesting and chick-rearing suggest sub-optimal habitat selection by Pied Avocets due to an incorrect assessment of the potential for successful reproduction of semi-natural sites, which may thus function as ecological traps.     

Estimating dispersal distributions at multiple scales: within‐colony and among‐colony dispersal rates,distances and directions in European Shags Phalacrocorax aristotelis

Knowledge of the rate, distance and direction of dispersal within and among breeding areas is required to understand and predict demographic and genetic connectivity and resulting population and evolutionary dynamics. However dispersal rates, and the full distributions of dispersal distances and directions, are rarely comprehensively estimated across all spatial scales relevant to wild populations. We used re‐sightings of European Shags Phalacrocorax aristotelis colour‐ringed as chicks on the Isle of May (IoM), UK, to quantify rates, distances and directions of dispersal from natal to subsequent breeding sites both within IoM (within‐colony dispersal) and across 27 other breeding colonies covering 1045 km of coastline (among‐colony dispersal). Additionally, we used non‐breeding season surveys covering 895 km of coastline to estimate breeding season detection probability and hence potential bias in estimated dispersal parameters. Within IoM, 99.6% of individuals dispersed between their natal and observed breeding nest‐site. The distribution of within‐colony dispersal distances was right‐skewed; mean distance was shorter than expected given random settlement within IoM, yet some individuals dispersed long distances within the colony. The distribution of within‐colony dispersal directions was non‐uniform but did not differ from expectation given the spatial arrangement of nest‐sites. However, 10% of all 460 colour‐ringed adults that were located breeding had dispersed to a different colony. The maximum observed dispersal distance (170 km) was much smaller than the maximum distance surveyed (690 km). The distribution of among‐colony dispersal distances was again right‐skewed. Among‐colony dispersal was directional, and differed from random expectation and from the distribution of within‐colony dispersal directions. Non‐breeding season surveys suggested that the probability of detecting a colour‐ringed adult at its breeding location was high in the northeastern UK (98%). Estimated dispersal rates and distributions were therefore robust to incomplete detection. Overall, these data demonstrate skewed and directionally divergent dispersal distributions across small (within‐colony) and large (among‐colony) scales, indicating that dispersal could create genetic and demographic connectivity within the study area.     

Safety first: terrestrial predators drive selection of highly specific nesting sites in colonial‐breeding birds

Nesting is a critical yet hazardous life stage for many birds. For colonial‐breeding birds, the conspicuousness of the colony to predators suggests immense pressure to select optimal colonial nesting sites. But what drives selection of those sites? As with solitary nesting birds, reducing access by predators may be the single most important factor. If so, knowledge of the predators involved and the attributes of different potential colony sites can allow us to predict the features that make a site especially safe. We examined the attributes of trees used by breeding colonies of metallic starlings Aplonis metallica in tropical Australia, and experimentally tested if those attributes prevented nest access by predatory snakes. Our surveys confirmed that tree choice by starling nesting colonies is highly non‐random, with all colonies located in tall trees in rainforest clearings, with no low branches and smooth bark. Experimental tests demonstrated that the climbing ability of predatory snakes depends upon bark rugosity, and that colony access by snakes depends on tree attributes such as bark rugosity and canopy connectivity. Our study confirms that colonial‐nesting starlings select colony sites that provide a safe refuge from predation. Intense predation pressure may have driven the evolution of stringent breeding habitat criteria in many other species of colonial‐breeding birds.     

Cooperative nest‐defence behaviour and territory quality in a resident and socially monogamous passerine

Nest defence is a fundamental aspect of parental care in secondary cavity‐nesting birds, and predation or competition for nesting sites can involve different defensive behaviours. Because habitat quality determines breeding success, we were interested in whether breeding pairs of the Eurasian nuthatch, Sitta europaea, established in more favourable environment also manifest higher probability of cooperative behaviour during their nest‐site defence. To explore this relationship, we quantified behavioural displays of both parents and analysed activity budget ethogram data from simulated territorial intrusions performed in the chick‐feeding phase with one conspecific and two different heterospecific stimuli (dummies of nuthatch, starling and woodpecker). We found that paired individuals shared their roles during nest‐site defence to a considerable extent. Males had a significantly higher number of attacks on intruders than females, and females performed more threat displays and controls of the brood than males. Multinomial analysis of the cooperative behaviour suggested that pairs in a high‐quality territory had higher probability of reciprocal substitution of different roles towards a balance between attacks, threat displays and nest controls. Contrary to this, pairs in a low‐quality territory had less likely pairwise combinations of simultaneous behavioural states that are associated with effective nest‐site defence. The difference in response probability according to territory quality was, however, highly variable in view of the stimulus that was used in simulated territorial intrusion. Because individual roles and the complex behavioural repertoire of pairs altered in response to territory quality and potential nest‐site competitor or brood predator, our results suggest that the cooperative nest‐defence behaviour could be linked to the breeding success of this year‐round territorial species living in a heterogeneous forest habitat.     

Intraspecific Food‐Robbing and Neighborhood Competition: Consequences for Anti‐Robber Vigilance and Colony Productivity

Most social animals have mechanisms to distinguish group members from outsiders, in part to prevent the exploitation of resources reserved for members of the group. Nevertheless, specialized thieves of the Neotropical ant, Ectatomma ruidum, also known as the ‘thieving ant’, regularly enter and steal resources from distinct, neighboring colonies. Here, we examine the mechanisms and consequences of thievery in a population of E. ruidum. We show that (1) individuals from nearby colonies were accepted more often than those from farther colonies; (2) rejection rates decreased as individuals interacted more with non‐nestmates from the same source colony; and (3) colonies that were experimentally treated to reduce thievery rates had improved productivity. The boost in productivity with thievery reduction was greater in low density populations than in high density populations. We conclude that, as in other species, thievery has negative fitness costs to E. ruidum. However, greater acceptance of neighbors than non‐neighbors and increased acceptance after habituation to non‐nestmates suggest a proximate explanation for the presence of thievery. Moreover, lower fitness costs of thievery at high nesting density, combined with observations of extraordinarily high densities of E. ruidum throughout its range, suggest there is little selection pressure among these ants to guard against thieves, thus providing an ultimate explanation why thievery persists among litter‐foraging ants.