Extended phenotypes and foraging restrictions: ant nest entrances and resource ingress in leaf‐cutting ants

Several factors may restrict the acquisition of food to below the levels predicted by the optimization theory. However, how the design of structures that animals build for foraging restricts the entry of food is less known. Using scaling relationships, we determined whether the design of the entrances of leaf‐cutting ant nests restricts resource input into the colony. We measured nests and foraging parameters in 25 nests of Atta cephalotes in a tropical rain forest. Ant flux was reduced to up to 60% at nest entrances. The width of all entrances per nest increased at similar rates as nest size, but the width of nest entrances increased with the width of its associated trail at rates below those expected by isometry. The fact that entrance widths grow slower than trail widths suggests that the enlargement of entrance holes does not reach the dimensions needed to avoid delays when foraging rates are high and loads are big. The enlargement of nest entrances appears to be restricted by the digging effort required to enlarge nest tunnels and by increments in the risk of inundation, predator/parasitoid attacks and microclimate imbalances inside the nest. The design of the extended phenotypes can also restrict the ingress of food into the organisms, offering additional evidence to better understand eventual controversies between empirical data and the foraging theory. Abstract in Spanish is available with online material.     

Intra-and intercolony patterns of nest dispersion in the ant Lasius neoniger: correlations with territoriality and foraging ecology

Colonies of the ant Lasius neoniger have multiple nest entrances that are distributed throughout a colony's foraging area. Associated with each nest entrance is a group of workers that show strong fidelity to that nest entrance. Territorial expansion, as indicated by increases in the number of nest entrances per colony, is correlated with foraging activity. Although there is variation between colonies in the seasonal pattern of territorial expansion, most nests become active in early summer, increase the size of the area foraged until midsummer, and then decrease the number of active nest entrances in late summer. Over the study plot as a whole, the dispersion pattern of nest entrances changed from clumped, or tending to be clumped, in early spring to random in mid-and late summer. Within colonies, nest entrances were significantly overdispersed. Intra-and interspecific competition negatively affected foraging, and workers from a given nest entrance were most successful at retrieving prey less than approximately 15–20 cm from the entrance. The average distance between nest entrances within a colony was 37.7±3.3 cm (mean±95% confidence interval, n=115), which is approximately twice the distance at which workers can retrieve prey. The polydomous nest structure of L. neoniger appears to partition territory within a colony by spatial subdivision of its foragers, and thus may reduce loss of prey to competitors.     

Size matters: nest colonization patterns for twig‐nesting ants

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The Importance of Where to Dump the Refuse: Seed Banks and Fine Roots in Nests of the Leaf‐Cutting Ants Atta cephalotes and A. colombica1

The location of the nutrient‐rich organic refuse produced by a leaf‐cutting ant colony varies among ant species. Atta cephalotes locate their organic refuse in subterranean chambers, whereas A. colombica place their organic refuse on the soil surface near the nest. We studied the effect of the absence or presence of external organic refuse on the abundance of fine roots and seed bank composition in the superficial horizons of ant nests. We sampled soils from ant nests or dumps and adjacent areas of 15 adult nests of A. cephalotes at La Selva (LS), Costa Rica, and of 15 of A. colombica nests on Barro Colorado Island (BCI), Panama. Soils from A. cephalotes nests did not differ from adjacent soils in abundance of fine‐root and seed diversity. In contrast, organic refuse from A. colombica nests was less diverse in seed composition (due to the great abundance of Miconia argentea) and had a greater abundance of fine roots than adjacent areas. Thus the external location of the ant‐nest organic refuse is potentially important in determining the different types of plant recolonization in abandoned or dead ant nests. The relative abundance of these Atta species may influence the structure and/or composition of tropical forests.     

Ant behaviour and seed morphology: a missing link of myrmecochory

Seed dispersal by ants (myrmecochory) is mediated by the presence of a lipid-rich appendage (elaiosome) on the seed that induces a variety of ants to collect the diaspores. When seeds mature or fall onto the ground, these ant species transport them to their nest. After eating the elaiosome, the seed is discarded in nest galleries or outside, in the midden or farther away, where seeds can potentially germinate. The final location of seeds with their elaiosomes removed was evaluated to assess the importance of possible handles (structures that ants can grasp to carry) in transporting ants during re-dispersal experiments of seeds from nests of six species of ants. The results indicate that seeds remained within the nest because the ants were not able to transport them out of the nest. As a consequence of the elaiosome being removed, small ant species could not take Euphorbia characias seeds out of their nests. Only large ant species could remove E. characias seeds from their nests. Attaching an artificial handle to E. characias seeds allowed small ant species to redistribute the seeds from their nests. On the other hand, Rhamnus alaternus seeds that have a natural handle after the elaiosome removal were removed from the nests by both groups of ant species. If a seed has an element that acts as a handle, it will eventually get taken out of the nest. The ants’ size and their mandible gap can determine the outcome of the interaction (i.e. the pattern of the final seed shadow) and as a consequence, could influence the events that take place after the dispersal process.     

Observation of plugging behaviour reveals entrance‐guarding schedule of morphologically specialized caste in Colobopsis nipponicus

The success of social insects is often attributed to the specialized morphologies and behaviours of workers. One of the most elaborate specializations in ant species is the workers’ plugging behaviour, in which their uniquely shaped heads are used for nest defence. These species generally nest in tree cavities. Without morphologically specialized workers (major workers) plugging the nest entrance, nests are easily attacked and replaced by conspecific and heterospecific cavity‐dwelling ants and other predators. Therefore, major workers are expected to defend the nest at all times. However, empirical evidence on the social time investment for nest defence is still lacking. Here, we investigated how major workers of Colobopsis nipponicus achieve nest defence (plugging behaviour) in terms of their work schedule. Our results showed that the nests had more major workers than entrances. The observation of artificial plastic nests over 6 days in the laboratory revealed that the nest entrance was guarded almost continuously by multiple major workers. The entrance‐guarding schedule was not equally divided among major workers, and there were no clear shift patterns in the timing of changeovers. We highlight the importance of investigating the time investment of workers to fully understand the defence strategies of this morphologically specialized caste.     

Complex Nesting Behavior by Two Neotropical Species of the Ant Genus Stenamma (Hymenoptera: Formicidae)1

Two Neotropical ant species, Stenamma expolitum Smith and S. alas new species (described here), exhibit three unusual nesting behaviors: (1) they build architecturally sophisticated nest entrances that elevate the nest opening away from the surface; (2) they maintain multiple identical nests but occupy only one of them; and (3) they keep a round “door pebble” at the nest entrance, with which they plug the opening in response to army ants. Adaptive hypotheses for these behaviors are discussed, including the possibility that there are multiple lines of defense against army ant predation.     

Intraspecific competition through food robbing in the harvester ant,Messor aciculatus (Fr. Smith), and its consequences on colony survival

Summary Intraspecific interference competition in the harvester ant,Messor aciculatus, was studied. Colonies of this species were found not to have territories. Some nests were located very close to each other, and the foraging areas of the neighbors usually overlapped. Even though the frequency with which alien and resident ants met was very high in the vicinity of the nest entrances, aggressive interactions between them rarely occurred. However, when hostile workers encountered each other, they exhibited a kind of ritualized combat and the winner ejected, but did not injure the loser. If any aliens entered the nest, some of them were pulled out, mainly by the residents.Aliens roaming near a neighbor's nest entrance ferociously attacked the residents carrying seeds in their mandibles and robbed them. On other occasions, aliens entered the nest and stole the collected seed. Although seed robbing and stealing occurred among neighboring colonies, there were remarkable differences in the frequency of their occurrence. The results of field observations and experiments suggest the existence of a dominance order among the neighbors. In one instance, extermination of an inferior colony by its neighbor was observed. The raider colony transferred the stored seeds from the nest of the inferior colony to its own and deposited the larvae and workers some distance away from the nest.The influence of ritualized combat and food robbing on colony activities, and the ecological significance of this interference behavior in terms of spatial distribution and temporal persistence of the nest sites, is discussed.     

Effects of the risk of competition and predation on large secondary cavity breeders

The effects of competition and risk of predation on secondary cavity breeders were examined between the 2008 and 2009 breeding seasons using an experimental design manipulating two nest entrance sizes (large entrances allowed Barn Owls (Tyto alba) to enter, while the small entrances excluded them). During the 2009 breeding season, the entrance sizes of nest boxes were exchanged, so that if during one year a nest box in a particular location had a small entrance, the second year it had a large entrance and vice versa. Barn Owls and Eurasian Kestrels (Falco tinnunculus) occupied 67.3 and 17.3%, respectively, of large entrance nest boxes. Significantly more Jackdaws (Corvus monedula), House Sparrows (Passer domesticus) and Scops Owls (Otus scops) bred in nest boxes with small than with large entrances. After nest box entrance sizes were exchanged, Barn Owls and smaller species did not breed in the same nest boxes with the new entrance size. Jackdaws probably did not breed in large entrance nest boxes due both to exploitation competition (Barn Owls and Eurasian Kestrels occupied the majority of large entrance nest boxes), and may also have avoided empty nest boxes because of the risk of interference competition; whereas smaller species may have also avoided large entrance nest boxes due to risk of predation.     

Geographical and interspecific variation and the nutrient‐enrichment hypothesis as an adaptive advantage of myrmecochory

In myrmecochory, the relocation of diaspores to ant nests may lead to the enhancement of plant fitness because ant nests and their middens are often richer in essential nutrients than surrounding areas. This idea is the basis of the nutrient‐enrichment hypothesis (NEH), which suggests that nutrient enrichment may be a major selective influence in the evolution of myrmecochory. However, there is little evidence regarding whether the greater plant performance and fitness enhancement in ant nests is due to nutrient enrichment or other benefits of directed dispersal. Here, we present the results of a large‐scale seed‐sowing experiment that tests the NEH in the ant‐dispersed perennial herb Helleborus foetidus, exploring geographical and inter‐ant taxa variation. Experiments were conducted in three well‐separated regions of the Iberian Peninsula, targeting the nests of major and minor local ant dispersers (nine ant species in total) and the soil beneath maternal plants as seed destinations. Seedling emergence, survival and early establishment rates, as well as variation in soil characteristics, were obtained for each seed destination at each region. Our results do not fully support the NEH in our study system. Instead, we found that the advantage of ant nest soil for establishment in H. foetidus was conditional. Differences in soil fertility and concomitant differences in seedling establishment between ant nests and beneath the canopy of maternal plants were observed in some regions and for some ant species, but not in others. Thus, the conditional outcomes arise from inconsistencies among regions, between stages of seedling regeneration and among ant species in the advantages of being dispersed to nests. Because variation in the guilds of ant dispersers of myrmecochore plants across their ranges is common, this study illustrates the need to consider geographic and inter‐ant taxa variation for a complete evaluation of the NEH.     

Secondary seed dispersal by ants in Neotropical cerrado savanna: species‐specific effects on seeds and seedlings of Siparuna guianensis (Siparunaceae)

1. Most woody plant species in tropical habitats are primarily vertebrate‐dispersed, but interactions between ants and fallen seeds and fruits are frequent. This study assesses the species‐specific services provided by ants to fallen arillate seeds of Siparuna guianensis, a primarily bird‐dispersed tree in cerrado savanna. The questions of which species interact with fallen seeds, their relative contribution (versus vertebrates) to seed removal, and the potential effects on seedling establishment are investigated. 2. Seeds are removed in similar quantities in caged and control treatments, suggesting that ants are the main dispersers on the ground. Five ant species attended seeds. Pheidole megacephala (≈0.4 cm) cooperatively transported seeds, whereas the smaller Pheidole sp. removed the seed aril on spot. Large (> 1.0 cm) Odontomachus chelifer, Pachycondyla striata, and Ectatomma edentatum individually carried seeds up to 4 m. Bits of aril are fed to larvae and intact seeds are discarded near the nest entrance. 3. Overall, greater numbers of seedlings were recorded near ant nests than in control plots without nests. This effect, however, was only detected near P. megacephala and P. striata nests, where soil penetrability was greater compared with controls. Soil nutrients did not differ between paired plots. 4. This study confirms the prevalence of ant–seed interactions in cerrado and shows that ant‐derived benefits are species‐specific. Ant services range from seed cleaning on the spot to seed displacement promoting non‐random spatial seedling recruitment. Although seed dispersal distances by ants are likely to be shorter than those by birds, our study of S. guianensis shows that fine‐scale ant‐induced seed movements may ultimately enhance plant regeneration in cerrado.     

THE FATE OF VIOLA SEEDS DISPERSED BY ANTS

Experimental plantings of the seeds of the two species Viola odorata and V. hirta were carried out to determine relative importance of three effects of myrmecochory: 1) clumping of seeds, 2) scarification of the testa and elaiosome removal, and 3) relocation into ant nests. The study site was a beech-larch wood in southern England. Data show that clumping slightly reduces seedling emergence, scarification and elaiosome removal slightly enhances it, while the nest environment significantly increases the rate of seedling emergence. Only the last effect is statistically significant. Seedlings from nests have larger first adult leaves. The possible impact of ant manipulation of seeds on seedling recruitment into myrmecochorous populations is discussed.     

The influence of oxygen stress on female choice for male nest structure in the common goby

This study examines the role of dissolved oxygen in modifying female mate choice in the common goby, Pomatoschistus microps. Experimental manipulations of nests showed that under normal (saturated) oxygen conditions, females preferred to spawn in nests with the most elaborate construction, that is, those with the most sand on top and the smallest entrance. Such manipulated nests proved less vulnerable to detection by an egg predator, the shore crab, Carcinus maenas, but these small entrances may reduce oxygenation of eggs. Indeed, in a low oxygen environment females with a choice between nests of high build and nests of low build did not significantly prefer either. This may have been due to weakened preference or perhaps detection of a mismatch between the male's parental quality and the increased ventilation needs of eggs when nest entrance sizes are small and ambient oxygen levels are low. When nests were not manipulated, those males that built small entrances fanned their eggs more often, such that hatching success in the absence of a predator was not related to the initial size of the nest entrance area. Thus, under normal oxygen conditions males may initially build nests with the smallest entrance they are capable of ventilating successfully, and females choosing such males gain from nest camouflage. Under low oxygen, the risk that manipulated males may be unable to compensate fully may outweigh such benefits, and females may use other criteria that signal willingness to provide parental care. Copyright 1999 The Association for the Study of Animal Behaviour.     

Nest orientation patterns in Salaria fluviatilis

The entrances of nests established under small stones by male river blennies Salaria fluviatilis in two rivers of the Ebro Basin in eastern Spain were randomly oriented in slow-flowing sites. In fast-flowing stretches or under large stones, however, nest entrances tended to open at an angle of c . 30° relative to current flow direction, i.e. near a south-east direction. As current velocity increased, males positioned their nest entrance closer and closer to the direction of flow. Selective nest entrance orientation reduced significantly the speed of current reaching the nest entrance such that current velocity was similar (5–7 cm s⁻¹) for all nests, regardless of stone size, prevailing current speed, or study site. Male mating success, measured as egg clutch area, however, was not related to current speed at the nest entrance but instead, it increased with nest stone size and decreased with deviations from a south-east direction. The reasons for female river blenny preference for this specific nest orientation are unknown but may be related to patterns of water flow, and hence oxygenation of the eggs, in the nest.     

Funnels,gas exchange and cliff jumping: natural history of the cliff dwelling ant Malagidris sofina

The Malagasy endemic ant Malagidris sofina (Bolton and Fisher 2014) nests on cliff faces in natural rock alcoves or clay banks. Colonies have single ergatoid queens and reproduce by fission. Each nest has a funnel-shaped entrance that projects horizontally from the cliff face. We examine three hypotheses for the function of the funnels—water exclusion, gas exchange and defense. Entrance funnels are relatively impermeable and divert water from nests, but simple tubes would achieve the same result. Consistent with the gas exchange hypothesis, projected funnel entrances likely increase gas exchange rates over sixfold compared to simple tubes and may increase air flow within the nest. Gas exchange may explain the recurrent evolution of funnel entrances in several ant lineages, especially among cliff dwelling species. We outline M. sofina defense responses to conspecifics and co-occurring ant species, and find no support for a defense role of entrance funnels. Workers display little aggression but respond to several species with an original form of nest defense––cliff jumping—in which workers drop off the cliff face while clinging to invaders and then return to their nest. M. sofina is a restricted range species under threat of extinction by habitat destruction. Its novel lifestyle underscores the urgency of exploration and conservation in a tropical biodiversity hotspot.     

Nest relocation in the slavemaking ants Formica subintegra and Formica pergandei: a response to host nest availability that increases raiding success

Social insects typically occupy spatially fixed nests which may thus constrain their mobility. Nevertheless, colony movements are a frequent component of the life cycle of many social insects, particularly ants. Nest relocation in ants may be driven by a variety of factors, including nest deterioration, seasonality, disturbances, changes in microclimate, and local depletion of resources. The colony movements of slavemaking ants have been noted anecdotally, and in recent studies such relocations were primarily attributed to nest deterioration or shifts to overwintering locations. In this study we explore nest relocations in large colonies of formicine slavemakers which occupy stable and persistent earthen nest mounds. We investigate the hypothesis that colony relocations of these slavemakers are best explained by efforts to improve raiding success by seeking areas of higher host availability. Five summers of monitoring the raiding behavior of 11–14 colonies of the slavemakers Formica subintegra and Formica pergandei revealed relatively frequent nest relocations: of 14 colonies that have been tracked for at least three of 5 years, all but one moved at least once by invading existing host nests. Movements tended to occur in the middle of the raiding season and were typically followed by continued raiding of nearby host colonies. Spatial patterns of movements suggest that their purpose is to gain access to more host colonies to raid: the distance moved is typically farther than the mean raiding distance before the move, which may indicate an effort to escape their local neighborhood. Furthermore, the mean distance of raids after relocation is shorter than the distance before relocation. For many slavemaking ant colonies, particularly those on the verge of relocating, raiding distance increased as the raiding season progressed. In addition, movements tended to be toward areas of higher local host density. Nest relocation is likely an important component of the ecology of slavemaking ants that contributes to the dynamic nature of their interaction with the host ant population.     

Removal of seeds from Neotropical frugivore droppings

In a Costa Rican rain forest, the majority of tree and shrub species have their seeds dispersed by vertebrates. Over a third of the species' seeds are of a size accessible to, and primarily carried away by, ants. Frugivorous bird droppings come in sizes from a few seeds to over a thousand, with number of seeds isometric with bird body mass. Dropping size decreases as seeds are scattered by rain. Larger droppings produced from fruits ofMiconia affinis (Melastomataceae) are discovered and recruited to more by ants, with a diverse guild of 22 ant species carrying away the seeds. Droppings with 64 and 16 seeds were used by a subset of the ant community exploiting 4-seed droppings, likely due to resource defense by aggressive species. Seeds in the smallest droppings stood the smallest chance of being removed. Although soil-nesting ants of the tribe Attini were the primary removers of seeds in this forest, a third of removal was by ants living in ephemeral litter nests. Seeds ofM. affinis were found in litter ant nests in 8 of 28 1-m² plots, suggesting that seed rain was not highly localized. Since litter nests are common, contain few seeds/nest and are often abandoned by their ants, litter ants may be the best candidates for ant-plant dispersal mutualisms.     

Hypoxic conditions and oxygen supply in nests of the mangrove ant, Camponotus anderseni, during and after inundation

The small ant Camponotus anderseni lives exclusively in twigs of the mangrove tree Sonneratia alba, and during inundation, the entrance hole is blocked with a soldier’s head which effectively prevents flooding. The nests can be very crowded, with the ants and coccids filling up to 50% of the volume, and due to their metabolic activity, the conditions in the nests during inundation become hypercapnic and hypoxic. Each nest has only one entrance, and the opening is quite small (1.56 ± 0.03 mm). The mean diameter of the galleries is 2.31 ± 0.23 mm, independent of the thickness of the twig and length of the nest. During normal conditions with open nests, the oxygen depletion is substantial in the part of the nest most distant from the opening, and in a 120 mm long nest the oxygen concentration can be as low as 15.7%. During simulated inundation, in which the nest entrances were blocked, the oxygen concentration dropped to very low levels (<0.5%) after one hour. After opening the nest entrance, the oxygen concentration increased again, but for a 100 mm long nest it took nearly 20 minutes before the concentration was back to the normal depressed level. Mathematical modelling of the steady-state oxygen concentrations in the innermost part of the nests shows a lower O₂ concentration than calculated. The time for equilibration of oxygen after inundation is longer than expected for small nests, presumably because the passive diffusion is obstructed by the nest contents. The “dilemma” faced by C. anderseni is to avoid drowning without suffering anoxia or hypercapnia, and they show a remarkable ability to adapt to the extreme conditions in the mangrove and exploit a niche where the density of other ants is insignificant. Received 13 December 2007; revised 30 July 2008; accepted 6 September 2008.     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Nest cavity orientation in black‐capped chickadees Poecile atricapillus: do the acoustic properties of cavities influence sound reception in the nest and extra‐pair matings?

Birds that nest in cavities may regulate nest microclimate by orienting their nest entrance relative to the sun or prevailing winds. Alternatively, birds may orient their nest entrance relative to conspecific individuals around them, especially if the acoustic properties of cavities permit nesting birds to better hear individuals in front of their nest. We measured the cavity entrance orientation of 132 nests and 234 excavations in a colour‐banded population of black‐capped chickadees Poecile atricapillus for which the reproductive behaviour of nesting females was known. Most chickadees excavated cavities in rotten birch Betula papyrifera, aspen Populus tremuloides and maple Acer saccharum. Nest cavities showed random compass orientation around 360° demonstrating that chickadees do not orient their cavities relative to the sun or prevailing winds. We also presented chickadees with nest boxes arranged in groups of four, oriented at 90° intervals around the same tree. Nests constructed in these nest box quartets also showed random compass orientation. To test the acoustic properties of nest cavities, we conducted a sound transmission experiment using a microphone mounted inside a chickadee nest. Re‐recorded songs demonstrate that chickadee nest cavities have directional acoustic properties; songs recorded with the cavity entrance oriented towards the loudspeaker were louder than songs recorded with the cavity entrance oriented away from the loudspeaker. Thus, female chickadees, who roost inside their nest cavity in the early morning during their fertile period, should be better able to hear males singing the dawn chorus in front of their nest cavity. Using GIS analyses we tested for angular‐angular correlation between actual nest cavity orientation and the azimuth from the nest tree to the territories and nest cavities of nearby males. In general, nest cavity entrances showed no angular‐angular correlation with neighbourhood territory features. However, among birds who followed a mixed reproductive strategy and nested in the soft wood of birch and aspen trees, nest cavity entrances were oriented towards their extra‐pair partners. We conclude that nest cavity orientation in birds may be influenced by both ecological and social factors.