Colony success of the submissive ant Formica fusca within territories of the dominant Formica polyctena

Abstract. 1. Twenty-three nests of the submissive ant Formica fusca L. were sampled in two adjacent territories of the dominant wood ant Formica polyctena Först. The nests were dug up at different distances from the wood-ant mounds. Distance is assumed to be inversely related to the extent of disturbance of F. fusca by F. polyctena.
2. Colony success of F. fusca was assayed by counting the numbers of workers (colony size), and worker and sexual offspring in a nest. For individual size, the head width and dry weight of fifty workers per nest were measured.
3. Colony size correlated positively with the numbers of worker and sexual offspring in one territory, but only with worker offspring in the other.
4. Distance from the wood-ant mound correlated positively with colony size and numbers of worker and female offspring in one territory. In the other territory distance correlated positively with number of females, but negatively with number of worker offspring. No significant correlations between distance and the size measurements were obtained.
5. Because of its nonaggressive behaviour F. fusca may nest fairly close to a wood-ant mound but is likely to show reduced abundances in terms of nest density and forager number, and, ultimately, lowered fitness.     

Nest structure and colony cycle of the Allegheny mound ant,Formica exsectoides Forel (Hymenoptera: Formicidae)

Summary The nest structure and colony cycle of a population of Allegheny mound ants,Formica exsectoides, were examined in central Michigan. The dispersion pattern of mounds was random. Nest structure and presence of brood were primarily determined by excavation of twenty-three nests over three intervals from June through September. Additional excavations of five nests in 1990 and ten nests in 1991 provided further details on nest structure and colony cycle. Most galleries occurred within the mound and upper 30 cm of soil, but some activity reached depths of 100 to 270 cm. Depth of nests showed little correlation with external measurements of height and diameter. Immature stages were recovered from two strata: the upper 20 cm of nest and mound and the lowest nest depths. Alate sexual forms were found in or near the mound in July, and numerous dealate queens were collected in September from peripheral galleries near the soil surface.     

Density- and size-dependent spacing of ant nests: evidence for intraspecific competition

Summary We present three lines of evidence which each suggest that intraspecific competition has significantly influenced the spacing patterns of Formica altipetens colonies. First, nearest-neighbor analysis of nest spacing patterns detected significant uniformity in six of eight plots. Second, there was a signifcant increase in the distance separating nearest neighbors as ant nest diameters increased. Third, ant nest density predicted substantial variation in the colony dispersion index, indicating the existence of a dispersion continuum at our study site.     

Brood raiding and the population dynamics of founding and incipient colonies of the fire ant, Solenopsis invicta

Abstract. 1. One of the first activities of minim workers in incipient fire ant nests is mutual brood raiding, the amalgamation of nests through the reciprocal stealing of brood and defection of workers.
2. Discrete mating flights created cohorts of incipient colonies. About 25% of founding nests survived the claustral period of 16–48 days (depending on season). Early incipient colony mortality was 5–6% per day. Over 60% of this mortality was accounted for through brood raiding. Most colonies raided when each cohort first became active, and many raided more than once. Raid size (number of nests, duration, distance) tended to increase during the summer as colonies grew through raiding. After the raiding period, nest mortality rate dropped 3–10-fold. Only 1–3% of founding nests were still alive at this time.
3. Queens from losing or failed nests tended to abandon their nests and attempted to enter successful ones, often following raiding trails to do so. This emigration was at least as successful as non-emigration in ultimately achieving the status of reproductrix of a successful nest (about 4%).
4. Brood raiding is a dominant process in early population dynamics, probably accounting for most of the early nest mortality. Its effect is to change the venue and unit of competition from nest-against-nest to a shifting aggregation of queens, workers and brood involving entire local populations. Nest thinning is thus very rapid, and the boost to the size of winning nests very large, allowing raiding colonies to win the competition for territory, and to achieve the early colony maturity so important to this, and other, weedy species. The importance of winning brood raids may also have driven increased minim production through the evolution of pleometrosis.     

Multiple breeders, breeder shifts and inclusive fitness returns in an ant

In social insects, colonies may contain multiple reproductively active queens. This leads to potential conflicts over the apportionment of brood maternity, especially with respect to the production of reproductive offspring. We investigated reproductive partitioning in offspring females (gynes) and workers in the ant Formica fusca, and combined this information with data on the genetic returns gained by workers. Our results provide the first evidence that differential reproductive partitioning among breeders can enhance the inclusive fitness returns for sterile individuals that tend non-descendant offspring. Two aspects of reproductive partitioning contribute to this outcome. First, significantly fewer mother queens contribute to gyne (new reproductive females) than to worker brood, such that relatedness increases from worker to gyne brood. Second, and more importantly, adult workers were significantly more related to the reproductive brood raised by the colony, than to the contemporary worker brood. Thus, the observed breeder shift leads to genetic benefits for the adult workers that tend the brood. Our results also have repercussions for genetic population analyses. Given the observed pattern of reproductive partitioning, estimates of effective population size based on worker and gyne samples are not interchangeable.     

Temporal and spatial variations of gyne production in the ant<Emphasis Type="Italic"> Formica exsecta</Emphasis>

Social insects have become a general model for tests of sex allocation theory. However, despite tremendous interest in the topic, we still know remarkably little about the factors that cause dramatic differences in sex allocation among local populations. A number of studies have suggested that environmental factors may influence sex allocation in ant populations. In polygynous (multiple queens per nest) populations of the ant Formica exsecta, sex allocation is extremely male biased at the population level, with only a small proportion of nests producing any gynes (female reproductive brood). We analysed the proportion of gyne-producing nests in 12 F. exsecta populations during three successive breeding seasons and found considerable temporal and spatial variability in the proportion of gyne-producing nests. The populations differed in a number of characteristics, including elevation, nest density, size of the nest mound, and number of nests per population. However, the proportion of gyne-producing nests was not associated with any of these geographic and demographic variables. Moreover, differences between populations in the production of gynes were not consistent between years. Thus, the proportion of gyne-producing nests appears to vary stochastically, perhaps because of stochastic variations in environmental factors. For example, year-to-year variations in the proportion of gyne-producing nests were associated with differences in spring weather conditions between years. The finding that gyne production varies greatly between years suggests that it may not always be adaptive at a local scale.     

A seasonal natural history of the ant, Odontomachus brunneus

A north Florida population of Odontomachus brunneus, a species of ponerine ants, was studied for a one-year period to determine the annual cycle of reproduction and colony growth, including the foraging biology and seasonal changes in nest architecture. The life cycle of O. brunneus is strongly seasonal. Colonies produce brood for 6 months and are broodless for 6 months. Alates are produced in mixed broods at the beginning of each season, consuming much of the colony’s energy reserves. These reserves recover slowly through foraging during the summer’s worker production, and rapidly after brood production ceases in October. The foraging population was estimated to average 77% (SD 22) of the workforce. This proportion was not related to colony size and female alates were also found to forage. Nest architecture was found to change seasonally, with winter nests being more than twice as deep as the average summer nest.     

An experimental analysis of joint nesting in the salamander Hemidaetylium scutatum (Caudata: Plethodontidae): the effects of population density

Three nesting behaviour patterns are documented in the plethodontid salamander Hemidactylium scutatum. A female may lay eggs (1) in a solitary nest and brood them, (2) in a joint nest and brood them as well as eggs of other females, or (3) in a joint nest that is brooded by another female. The hypothesis that population density was positively associated with joint nesting was tested by following two populations for 5 years and by experimentally manipulating the population density of nesting females in artificial habitats for the latter 2 years. The proportion of joint nests did not vary with density, although joint nests tended to contain eggs of more females at the high population density. Joint nests were usually brooded by one female; thus, most females that laid eggs in joint nests did not brood them at high density. The reproductive success, as measured by survival of embryos, of solitary and joint nesters was equivalent. Joint nests were deserted less often, however, which decreased the probability of catastrophic mortality. The number of days of brooding was significantly positively correlated with loss of body mass of females, suggesting a cost to brooding behaviour. Joint nesting with solitary brooding is not explained by aggressive usurpation of nests or by brood parasitism.     

Spatial patterns, temporal variability, and the role of multi-nest colonies in a monogynous Spanish desert ant

Abstract 1. The colonies of the Spanish desert ant Cataglyphis iberica are polydomous. This study describes the temporal and spatial patterns of the polydomy in this species at two different sites, and presents analyses of its role in reducing the attacks of the queen over sexual brood, and in allowing better habitat exploitation.
2. The spatial distribution of nests was clumped while colonies were distributed randomly. Mean nearest neighbour distance ranged from 3.4 to 7.0 m for nests and from 12.3 to 14.1 m for colonies. Distance of foragers searching for food varied among nests: mean values were between 6.1 and 12.6 m.
3. At both sites, the maximum number of nests per colony occurred in summer, during the maximum activity period of the species. Colonies regrouped at the end of this period but overwintered in several nests.
4. Nest renewal in C. iberica colonies was high and showed great temporal variability: nests changed (open, close, re-open) continuously through the activity season and/or among years. The lifetime of up to 55% of nests was only 1–3 months.
5. Polydomy in C. iberica might decrease the interactions between the queen and the sexual brood. In all colonies excavated just before the mating period, the nest containing the queen did not contain any virgin female. Females were in the queenless nests of the colony.
6. The results also suggest that polydomous C. iberica colonies may enhance habitat exploitation because foraging activity per colony increases with nest number. The relationship between total prey input and foraging efficiency and number of nests per colony attains a plateau or even decreases after a certain colony size (four to six nests). This value agrees with the observed mean number of nests per colony in C. iberica .     

Colony size and nest density predict the likelihood of parasitism in the colonial Southern Red Bishop Euplectes orix– Diderick Cuckoo Chrysococcyx caprius system

Colonial nesters are potentially susceptible to brood parasitism because they present an aggregated source of accessible nests to brood parasites. However, colonial breeding may confer fitness advantages on potential host species that outweigh the costs of parasitism, particularly if colonies are large or dense enough for corporate vigilance to deter parasitism. In addition, some studies have suggested that the spatial habitat structure near host populations is a critical determinant of parasitism rate, with cuckoos targeting those colonies close to cover. We examined the determinants of the likelihood of parasitism by Diderick Cuckoos Chrysococcyx caprius on their main southern African host, the colonially breeding Southern Red Bishop Euplectes orix , from 24 colonies and 1141 nests over two seasons. The likelihood of parasitism by Diderick Cuckoos decreased significantly with increasing colony size and nest density, but not with distance to cover from a colony, providing no support for the spatial habitat structure hypothesis. We suggest that proximate constraints, such as visibility in the semi-closed nest, limit the ability of the host to recognize the Cuckoo egg, giving rise to low rates of rejection of Cuckoo eggs. However, the Cuckoo cannot exploit this apparent weakness when the host breeding population is in large colonies that are protected by corporate vigilance. Consequently, the Diderick Cuckoo – Southern Red Bishop system may represent an evolutionary equilibrium in host and parasite defences.     

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.     

The economics of brood raiding and nest consolidation during ant colony founding

Summary The most dangerous time for an ant colony is during the founding stage when the small colony is vulnerable to predation and competition. Colonies can grow more rapidly when multiple queens cooperate in raising the first worker brood (pleometrosis) or by raiding other incipient colonies for their brood. This brood raiding has been proposed to be the primary force selecting for pleometrosis, i.e. multiple-queen colonies may have a considerable advantage in destroying neighbours by aggressively stealing their brood. An alternative hypothesis is that incipient nests are part of a larger, interconnected population structure and that brood raiding reflects cooperative pleometrosis with subdivided colonies. A simple mathematical model supports the second hypothesis: workers of incipient colonies are especially favoured to peaceably abandon their nest and join with other colonies if the queens are related or queens from raided colonies can infiltrate the raiding colony. The latter condition is often met in ant species that brood raid and particularly exemplified in fire ants (Solenopsis invicta), where brood raiding involves little mortal combat and combines with pleometrosis to rapidly increase colony size. It is proposed that the term nest consolidation should replace brood raiding to more accurately reflect the relatively non-aggressive and potentially apparently cooperative nature of interactions between incipient ant colonies.     

Experimental enlargement of nest size does not increase risk of predation or brood parasitism in the Great Reed Warbler Acrocephalus arundinaceus

We assessed whether nest size affects the probability of nest loss using dyads of large and small (large being twice the size of small) inactive Great Reed Warbler Acrocephalus arundinaceus nests placed at similar sites in Great Reed Warbler territories. Large nests were not predated significantly more frequently than small nests. Experimentally enlarged active Great Reed Warbler nests suffered non‐significantly higher predation compared with non‐manipulated control nests. Our experiments did not support the nest‐size hypothesis and suggested that nest size does not appear to be a factor affecting the risk of nest predation in this species. The probability of brood parasitism by the Common Cuckoo Cuculus canorus was also unaffected by experimental nest enlargement, supporting the commonly accepted hypothesis that the Common Cuckoo searches for suitable host nests by host activity during nest building rather than nest size.     

Worker populations of Formica lugubris Zett. nests in Irish plantation woods

Abstract. 1. Estimates of worker populations of Formica lugubris Zett. nests in Irish plantation woods were obtained using capture—mark—recapture methods. Forager numbers were estimated by paint-marking individuals on the foraging routes and 'recapturing' by traffic counts. Colony-size was estimated using Stradling's (1970) ³²P-radiolabelling technique. Worker brood was estimated by excavation and direct count.
2. Forager numbers ranged from 6906 to 64 686 (eleven nests), colony-size ranged from 9797 to 71 052 (five nests) and worker brood ranged from 9809 to 16 269 (four nests).
3. Forager number was highly correlated with the average traffic on individual routes and the nest forager populations declined rapidly in >30-year-old nests.     

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.     

Barn swallows prefer to nest at sites hidden from neighboring nests within a loose colony

Some loosely colonial species scatter their nests within a colony. Most studies on mechanisms that generate such scattered nests within a colony have primarily focused on the measurement of distance related to the intensity of interactions among conspecific neighbors. However, these interactions can change not only with distance between nests but also with other factors. In particular, whether the nest is hidden from neighbors can strongly affect the intensity of interaction for birds that usually use sight for communication. To show the possible benefit of nesting at sites hidden from neighboring nests, we studied the nest arrangements and settlement patterns within a barn swallow colony. We tested whether the swallows prefer nesting at sites hidden from neighboring nests in cattle barns that are often used as colony sites by the swallows in Japan. We found that the sides of ceiling beams hidden from neighbors were preferred as nesting sites. A randomization test suggested that the rectangular sections occupied by swallows were not spaced out within a colony. The results suggested that the swallows preferred nesting sites hidden from their neighbors and did not space out their nests within a colony. This paper also refutes intraspecific brood parasitism as a plausible function of this behavior.     

Responses of naive yellow warblers to a novel nest predator

Naive parental yellow warblers, Dendroica petechia, were tested with a mounted eastern grey squirrel, Sciurus carolinensis, a novel predator on the study site, during the nest-building, egg-laying, incubation and nestling stages of the nesting cycle. Testing parents only once avoided the possibility that the nest defence behaviour of the parents was modified by reinforcement and loss of fear. In 96% of 80 trials, the female parent responded first to the model. Males rarely showed nest defence behaviour. Female nest defence as measured by defensive vocalizations (i.e., ‘chipping’), closest and modal distance of approach to the model, distraction displays, strikes, and close passes or hovers, did not increase significantly over the breeding season. ‘Chipping’ frequency by both sexes tended to increase after the incubation period but not significantly so. Distraction displays increased significantly only between nest-building and nestling stages. Age, clutch or brood size, and nest success were not correlated with the intensity of nest defence. Females defending exposed nests gave more distraction displays than those defending cryptic nests.     

Effectiveness of nest defence in the Acadian Flycatcher Empidonax virescens

We used presentations of models to determine the effectiveness of nest defence in the Acadian Flycatcher Empidonax virescens against a nest predator (Blue Jay Cyanocitta cristata ) and a brood parasite (Brown-headed Cowbird Molothrus ater ). Principal components analysis (PCA) of four component variables of nest defence (call rate, swoop rate, closest approach and number of adults) generated a measure of overall nest defence (aggression). We determined effectiveness of defence by looking for correlations between measures of defence and measures of nest success (nest predation and brood parasitism). We also determined whether nest defence increased with clutch size, nestling age and time in the breeding season. Defence against model Brown-headed Cowbirds did not correlate with levels of parasitism, clutch size, age of young or time of breeding. There was, however, a strong, but insignificant, trend for nests with high levels of all measures of defence to suffer less from brood parasitism. Aggression, vocalization rate, closest approach and number of adults defending against models of predatory Blue Jays correlated positively with nesting success during the egg stage but not the nestling stage of the nesting cycle. Aggression, vocalization rate, closest approach correlated with clutch size and age of the brood. These results suggest that nest defence can effectively deter nest predators, but may be less effective against brood parasites. Different behavioural components of nest defence may work at different stages of the nest cycle and against different nest predators. The components of nest defence that correlated with nest success also correlated with clutch value, a result consistent with hypotheses on the evolution of nest defence.     

Feeding frequency influences crèching age in the Dalmatian pelican, Pelecanus crispus

Crèching behaviour is common in colonial seabirds; nevertheless, the factors inducing chicks to aggregate remain relatively poorly understood. It has been proposed that brood size, laying date and nest attendance are important factors in the formation of a crèche. Moreover, in most species of pelicans, chicks join crèches following the development of homoeothermy and coincident with the end of the brooding behaviour. We studied effects of feeding rate, nest attendance, brood size, laying date and homoeothermy on the age at which chicks entered the crèche at a colony of Dalmatian pelicans (Pelecanus crispus), in Srebarna, Bulgaria. Single chicks were fed more frequently than chicks from two-chick broods. Unlike American white pelicans (Pelecanus erythrorhynchos), Dalmatian pelicans maintained brooding behaviour a further 9 days after chicks had developed thermoregulation abilities. In contrast to nests with two chicks, nests with only one chick were never left unattended by the parents before the chick reached the crèching stage. Laying date, nest attendance and brood size did not affect the age that the chick entered the crèche. The age the chick entered the crèche was not correlated with the age of homoeothermy acquisition, but chicks significantly joined the crèche at younger ages when the mean number of feeds per chick per day during the rearing period in the nest was higher. This result suggests an implication of growth rate in the crèching age. Joining the crèche earlier can provide benefits that could have strong implications for the chicks’ future reproductive lives.     

Density-dependent nest predation in waterfowl: the relative importance of nest density versus nest dispersion

When nest predation levels are very high or very low, the absolute range of observable nest success is constrained (a floor/ceiling effect), and it may be more difficult to detect density-dependent nest predation. Density-dependent nest predation may be more detectable in years with moderate predation rates, simply because there can be a greater absolute difference in nest success between sites. To test this, we replicated a predation experiment 10 years after the original study, using both natural and artificial nests, comparing a year when overall rates of nest predation were high (2000) to a year with moderate nest predation (2010). We found no evidence for density-dependent predation on artificial nests in either year, indicating that nest predation is not density-dependent at the spatial scale of our experimental replicates (1-ha patches). Using nearest-neighbor distances as a measure of nest dispersion, we also found little evidence for "dispersion-dependent" predation on artificial nests. However, when we tested for dispersion-dependent predation using natural nests, we found that nest survival increased with shorter nearest-neighbor distances, and that neighboring nests were more likely to share the same nest fate than non-adjacent nests. Thus, at small spatial scales, density-dependence appears to operate in the opposite direction as predicted: closer nearest neighbors are more likely to be successful. We suggest that local nest dispersion, rather than larger-scale measures of nest density per se, may play a more important role in density-dependent nest predation.