Prey spectra of two swarm-raiding army ant species in East Africa

Although swarm-raiding army ants are considered keystone predators in tropical forest ecosystems, information on the prey spectra of most species is based on anecdotal reports and not on systematic studies with extensive sampling. We analysed prey samples of 18 colonies of the two afrotropical species Dorylus ( Anomma ) molestus and Dorylus ( Anomma ) wilverthi (4289 prey items in total) to examine the prey composition variation within and between species and to determine the best methodology to obtain reliable prey spectrum estimates for a given species, site and season. Variation in prey composition was substantial for D. molestus even within a single site and season, with the biomass proportion of the most important prey type differing by a factor of 12. Conclusions from studies using small samples sizes may thus be misleading. We demonstrate that the method of assessing prey spectra in terms of relative prey item numbers often produces biased results, and therefore recommend relative prey biomass as the more useful parameter. The near absence of earthworms, which always constituted a substantial part of the D. molestus prey, in the diet of D. wilverthi is interpreted to result from subtle differences in swarm-raiding behaviour between these two species, but could alternatively also be due to low availability. Similar studies recording prey composition as biomass proportions and analysing large samples sizes from many colonies are needed to understand the effect of army ant swarm raids on invertebrate communities in afrotropical forests.     

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 .     

The ontogeny of foragwehaviour in desert ants, Cataglyphis bicolor

Abstract. 1. Individually foraging desert ants, Cataglyphis bicolor , exhibit short foraging lives (half lifetime, i.e. half-time of the exponential decay function: 4.5 days), in which they perform 3.7 ± 1.9 foraging runs per day.
2. During their short lifetime foraging period the ants increase the duration of their foraging round trips (up to 40.0 ± 24.6 min per run), the maximal distance of individual foraging runs (up to 28.2 ± 4.1 m), and their foraging success, i.e. the ratio of successful runs to the total number of runs (up to 0.70).
3. The parameter that increases most dramatically during a forager's lifetime is direction fidelity, i.e. the tendency to remain faithful to a particular foraging direction.
4. A model based on some simple behavioural rules is used to describe the experimental findings that within an isotropic food environment individual ants develop spatial foraging idiosyncrasies, and do so at a rate that increases with the food densities they encounter.
5. Finally, it is argued that in functional terms direction fidelity is related to the navigational benefits resulting from exploiting familiar (route-based) landmark information, and hence reduces round-trip time and by this physiological stress and predatory risk.     

Seasonal spatial dynamics and causes of nest movement in colonies of the invasive Argentine ant (Linepithema humile)

Abstract.  1. Colony organisation and movement behaviour of the Argentine ant ( Linepithema humile ) was studied over 3 years in field populations in California and in captive colonies in the laboratory. This invasive species is highly polydomous and unicolonial; colonies consist of expansive and fluid networks of nests and trails. The spatial and temporal organisation of colonies may contribute to ecological dominance.
2. Argentine ant nests and inter-nest trails shift in size, abundance, and location, so that colony networks are spatially contracted in the winter and expanded spring to autumn. Colonies occupy permanent sites; ants migrated to and from the same winter nest locations year after year, and occupied 30% of the same nests repeatedly during seasonal migrations.
3. Nests were moved on average 2–3 m. Forty-two per cent were occupied less than 1 month, 4% the entire study, and the other 54% lasted 3.9 ± 2.3 months (mean ± SD).
4. Nests were located within 2–4 m of woody plants, in warm sites in the winter and cool sites in the summer. Both humidity and food availability influenced nest-site choice in laboratory colonies. However, when faced with a trade-off between factors, the ants chose humid nest boxes over nest boxes near food, and ants moved nests only in response to changes in humidity and not distance to food.
5. The results indicate that L. humile colonies are seasonally polydomous, and that nest movements are driven by changes in microclimate. Colony organisation maintains high local density and increases food supply, which may improve the competitive ability of L. humile colonies and reduce opportunities for species coexistence.     

Male parentage in army ants

In most social insects workers do not mate, but have retained the ability to produce haploid eggs that can develop into viable male offspring. Under what circumstances this reproductive potential is realized and how the ensuing worker-queen conflict over male production is resolved, is an area of active research in insect sociobiology. Here we present microsatellite data for 176 males from eight colonies of the African army ant Dorylus (Anomma) molestus. Comparison with worker genotypes and inferred queen genotypes from the same colonies show that workers do not or at best very rarely reproduce in the presence of the queen. Queens of D. (A.) molestus are known to be highly multiply mated. This implies that workers are on average more closely related to queen sons than to other workers' sons, so that our results are consistent with predictions from inclusive fitness theory. It remains unknown, however, whether worker sterility is maintained by active worker policing or by self-restraint.     

Foraging trip duration of bumblebees in relation to landscape-wide resource availability

Abstract.  1. The study tested the hypotheses that bumblebees have shorter foraging trips in environments that provide abundant resources than in environments that provide sparse resources, and that shorter foraging trips translate into greater colony growth.
2. Six even-aged Bombus terrestris colonies were established in contrasting resource environments. Three colonies had access to abundant resources ( Phacelia tanacetifolia fields with high flower densities), and three colonies were placed in an environment with sparse resources (scattered semi-natural habitats with food plants at lower densities).
3. A total of 870 foraging trips of 220 marked B. terrestris foragers were observed using automated camcorder recordings.
4. The duration of foraging trips was shorter in environments with abundant resources (66 ± 4.6 min) than in environments with sparse resources (82 ± 3.7 min). Within 34 days colonies that had access to abundant resources gained significantly more weight (129 ± 40 g) than colonies foraging on sparse resources (19 ± 7 g).
5. Thus, the spatial distribution and quality of resources at landscape level affected the duration of foraging trips and the colony growth. It was concluded that future conservation schemes need to improve the spatial and temporal availability of resources in agricultural landscapes to counteract the ongoing decline of bumblebees.     

Species and site differences in Neotropical army ant emigration behaviour

Abstract 1. Along with large wingless queens and group foraging raids, periodic colony movements or emigrations are a defining feature of army ant behaviour. Emigrations are important because they regulate spacing among colonies, and provide access to new foraging sites. 2. Raid and emigration activity of army ant communities was measured at four Neotropical sites using a standardised sampling protocol. Sampling of emigration activity was carried out throughout the diel cycle. 3. Both raid and emigration activity occurred throughout the day and night. Emigration activity was highest under cover of darkness, but sites and species differed in the diel timing of emigrations. These site and species differences have implications for the evolution of emigration behaviour, as well as for the community ecology of army ants.     

Driver Ants Invading a Termite Nest: Why Do the Most Catholic Predators of All Seldom Take This Abundant Prey?

Driver ants ( i.e. , epigaeic species in the army ant genus Dorylus , subgenus Anomma ) are among the most extreme polyphagous predators, but termites appear to be conspicuously absent from their prey spectrum and attacks by driver ants on termite nests have not yet been described. Here, we report a Dorylus ( Anomma ) rubellus attack on a colony of the fungus-growing termite Macrotermes subhyalinus that was observed during the dry season in a savannah habitat in Nigeria's Gashaka National Park. It was estimated that several hundred thousand termites (probably more than 2.4 kg dry mass) were retrieved. The apparent rarity of driver ant predation on Macrotermes nests may be explained by different habitat requirements, by the fact that these ants mostly forage aboveground, by efficient termite defense behavior and nest architecture that make entry into the nest difficult, and finally by driver ant worker morphology, which differs remarkably from that of subterranean Dorylus species that regularly invade and destroy termite colonies.     

Phenetic studies of African army ant queens of the genus Dorylus (Hymenoptera: Formicidae)

Multivariate analyses of queens of fourteen species of the Old World army ant genus Dorylus were undertaken to assist in clarifying the status of the six included subgenera. Results were compared with two similar studies based on major workers and males, respectively. Queens are phenetically less divergent than either of the other two phena examined, although similar sub-generic relationships were observed. Taken together, these phenetic studies suggest that the subgenera Rhogmus and Alaopone are deserving of continued individual status, but that species of Anomma and Dorylus (s.s.) form one diverse taxon. A conservative classification would place Dichthadia and Typhlopone species together with the composite Dorylus (s.s.) - Anomma taxon.     

On a wing and a prayer: the foraging ecology of breeding Cape cormorants

The Cape cormorant Phalacrocorax capensis is unusual among cormorants in using aerial searching to locate patchily distributed pelagic schooling fish. It feeds up to 80 km offshore, often roosts at sea during the day and retains more air in its plumage and is more buoyant than most other cormorants. Despite these adaptations to its pelagic lifestyle, little is known of its foraging ecology. We measured the activity budget and diving ecology of breeding Cape cormorants. All foraging took place during the day, with 3.6 ± 1.3 foraging trips per day, each lasting 85 ± 60 min and comprising 61 ± 53 dives. Dives lasted 21.2 ± 13.9 s (maximum 70 s), attaining an average depth of 10.2 ± 6.7 m (maximum 34 m), but variability in dive depth both within and between foraging trips was considerable. The within-bout variation in dive depth was greater when making shallow dives, suggesting that pelagic prey were targeted mainly when diving to <10 m. Diving ecology and total foraging time were similar to other cormorants, but the time spent flying (122 ± 51 min day⁻¹, 14% of daylight) was greater and more variable than other species. Searching flights lasted up to 1 h, and birds made numerous short flights during foraging bouts, presumably following fast-moving schools of pelagic prey. Compared with the other main seabird predators of pelagic fish in the Benguela region, Cape gannets Morus capensis and African penguins Spheniscus demersus , Cape cormorants made shorter, more frequent foraging trips. Their foraging range while feeding small chicks was 7 ± 6 km (maximum 40 km), similar to penguins (10–20 km), but less than gannets (50–200 km). Successful breeding by large colonies depends on the reliable occurrence of pelagic fish schools within this foraging range.     

New role for majors in Atta leafcutter ants

Abstract.  1.  Atta (Hymenoptera: Formicidae) leafcutter ants display the most polymorphic worker caste system in ants, with different sizes specialising in different tasks. The largest workers (majors) have large, powerful mandibles and are mainly associated with colony defence.
2. Majors were observed cutting fallen fruit and this phenomenon was investigated in the field by placing mango fruit near natural Atta laevigata and Atta sexdens colonies in São Paulo State, Brazil.
3. Ants cutting the fruit were significantly heavier (mean = 49.1 mg, SD = 11.1 mg, n = 90) than the ants carrying the fruit back to the nest (mean = 20.9 mg, SD = 9.2 mg, n = 90).
4. Fruit pieces cut by majors were small (mean = 15.9 mg), approximately half the weight of leaf pieces (mean = 28.5 mg) cut and carried by media foragers. It is hypothesised that it is more difficult to cut large pieces from three-dimensional objects, like fruit, compared to two-dimensional objects, like leaves, and that majors, with their longer mandibles, can cut fruit into larger pieces than medias.
5. The study shows both a new role for Atta majors in foraging and a new example of task partitioning in the organisation of foraging.     

Low intraspecific aggression level in the polydomous and facultative polygynous ant Ectatomma tuberculatum

Nestmate recognition is a key feature of social insects, as it preserves colony integrity. However, discrimination of non‐nestmates and nestmate recognition mechanisms are highly variable according to species and social systems. Here, we investigated the intraspecific level of aggression in the facultative polygynous and polydomous ant, Ectatomma tuberculatum Olivier (Hymenoptera: Formicidae: Ectatomminae), in a population with a strong genetic structure. We found that the intraspecific level of aggression was generally low in this population of E. tuberculatum. However, the level of aggression was significantly correlated with the geographical distance, suggesting that both genetic and environmental cues could be involved in nestmate recognition and discrimination mechanisms. Moreover, polydomy was confirmed by the absence of aggression between workers from nests at a distance of 3 m, while the level of aggression was significantly higher between workers from nests separated by a distance of 10 or 1300 m. Field experiments showed that the low level of aggression between neighbouring colonies was associated with shared foraging areas, which could suggest that familiarization processes may occur in this species. We propose that the particular social organization of this species, with secondary polygyny, polydomy, and budding, may have favoured a high acceptance threshold, because of the low probability of interactions with unrelated conspecifics, the high cost of erroneously rejecting nestmates, and the low cost of accepting non‐nestmate workers. The resulting open recognition system can thus allow privileged relationships between neighbouring colonies and promote the ecological dominance of E. tuberculatum in the mosaic of arboreal ants.     

Dispersal and foraging behaviour of Platygaster californica: hosts can't run, but they can hide

Abstract.  1. Host–parasitoid models often identify foraging behaviour and dispersal distance as important for system persistence.
2. Laboratory observations and field trials were used to characterise foraging behaviour and dispersal capability of Platygaster californica Ashmead (Platygasteridae), a parasitoid of the gall midge Rhopalomyia californica Felt (Cecidomyiidae).
3. Although foraging parasitoids meticulously searched plants in laboratory observations, none of the laboratory trials resulted in 100% parasitism, and the proportion of parasitism declined as midge egg density increased.
4. The field trials showed that the distribution of parasitism over distance from a central release point was hump-shaped, as predicted by a simple diffusion model. Mean parasitoid dispersal distance was 4.5 m, considerably farther than the 1.7 m mean midge dispersal found in previous work.
5. Although the parasitoid appears to search thoroughly for midge eggs and to disperse farther than the midge, the results of this study show how this host–parasitoid system may persist due to spatially variable incomplete parasitism.     

The foraging ecology of the army ant Eciton rapax: an ergonomic enigma?

Abstract. 1. The army ant Eciton rapax (F. Smith) produces longer raid systems than any other member of its genus and it is a specialist predator of forest floor and understory ants such as species of Camponotus, Odontomachus and Pachycondyla.
2. Allometrical analysis confirms that E. rapax is the only member of its genus without distinct physical castes among its workers: its foraging population is entirely monomorphic and there are no majors.
3. The workers of E. rapax are distributed over a considerable size range, and there are distinct divisions of labour within these colonies: small workers tend to stay in the nests and among the larger foraging workers those retrieving prey items are significantly bigger than the rest.
4. An analysis of foraging efficiency and worker performance in E. rapax suggests that transport costs, resulting from the great distances that workers travel during raids and emigrations, are one of the selection pressures that have favoured the evolution of large monomorphic workers in this species.     

Vigilance during Preening in Coots Fulica atra

In birds, vigilance during feeding is usually linked with (1) the many‐eyes‐hypothesis, (2) the dilution effect, and (3) the intraspecific scramble competition hypothesis. To exclude competition for resources as a driving force in vigilance research into activities other than feeding is necessary. Additionally, variables such as nearest neighbour distance and position within a flock are supposed to influence vigilance and vigilance of singletons should be compared with vigilance of flock members. I studied vigilance during preening in Eurasian Coots (Fulica atra) and counted the number of scans per minute. A total number of 117 coots were sampled with 16 of them preening alone. Coots preening alone showed a significantly higher scan rate. I found a significant negative correlation between vigilance and flock size while nearest neighbour distance correlated positively with vigilance. Further, vigilance was higher in individuals at the edge. A general linear model using nearest neighbour distance and flock size within 10 m as covariates also revealed a significant influence of flock size but not of nearest neighbour distance. The centre‐edge effect still remained significant. These results indicate that flock size is the most influential predictor of vigilance in preening coots, followed by spatial organization. However, using nearest neighbour distance instead of flock size also produced a significant model as did the flock size measurement within a radius of 20 m. As scramble competition could be ruled out, the flock size effect may be indeed related to predation.     

Multi-scale pattern analysis of a mound-building termite species

Termite mounds are a widespread feature in most African savannas. These structures exhibit high nutrient contents and often host a special vegetation composition. In this study, we analysed mound distribution patterns of a fungus-growing termite species, Macrotermes michaelseni, an important ecosystem engineer in the savannas of Namibia. Inhabited mounds taller than 0.7 m were regularly distributed. We view this pattern as a result of intraspecific competition. The heights of mounds taller than 0.7 m were correlated positively with their distance, such that mounds closer together, i.e. up to inter-mound distances of approximately 50 m, tended to be smaller than average. This indicates that intraspecific competition for foraging areas controls mound distribution pattern and colony size. Differences between mound heights increased on the spatial scale up to inter-mound distances of 80 m. We assume that the foundation of new colonies is only possible in unoccupied patches. In such patches, young colonies are able to occur close together as they have a relatively low foraging demand and therefore a low spatial demand. In contrast, their critical distance to taller colonies with higher foraging demands is rather large, which leads to the observed increasing difference of mound heights with increasing distances between them.     

Intraspecific aggression, colony identity and foraging distances in Sudanese Microtermes spp. (Isoptera: Termitidae: Macrotermitinae)

Abstract. 1. Intraspecific aggression between termite major workers was used to obtain estimates of foraging distances for three Microtermes species in Sudan.
2. Maximum foraging distance recorded for M. sp. nr albopartitus (Sjöstedt) in Khartoum was 11.3 m, giving an estimated minimum colony area of 100 m². This is probably an underestimate. In the Tokar Delta, individuals from single colonies of M. najdensis Harris were encountered up to 42 m apart, giving a colony area of 1390 m².
3. Intraspecific aggression could not be used reliably to distinguish members of different colonies of M. lepidus Sjöstedt. Soil barrier formation between groups of workers in petri dishes may be of use as a supplementary technique, especially for species showing no clear aggression.
4. Experiments on isolated laboratory colonics of M. sp. Nr lepidus Sjöstedt, established from alates collected in Khartoum, further confirmed the value of inter-colony aggression for indicating colony identity.
5. The complications introduced into estimation of subterranean termite foraging areas by overlap and interdigitation of colonies are discussed.     

Both female castes contribute to colony emigration in the polygynous ant Mystrium oberthueri

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Extreme queen-mating frequency and colony fission in African army ants

Army ants have long been suspected to represent an independent origin of multiple queen-mating in the social Hymenoptera. Using microsatellite markers, we show that queens of the African army ant Dorylus (Anomma) molestus have the highest absolute (17.3) and effective (17.5) queen-mating frequencies reported so far for ants. This confirms that obligate multiple queen-mating in social insects is associated with large colony size and advanced social organization, but also raises several novel questions. First, these high estimates place army ants in the range of mating frequencies of honeybees, which have so far been regarded as odd exceptions within the social Hymenoptera. Army ants and honeybees are fundamentally different in morphology and life history, but are the only social insects known that combine obligate multiple mating with reproduction by colony fission and extremely male-biased sex ratios. This implies that the very high numbers of matings in both groups may be due partly to the relatively low costs of additional matings. Second, we were able to trace recent events of colony fission in four of the investigated colonies, where the genotypes of the two queens were only compatible with a mother-daughter relationship. A direct comparison of male production between colonies with offspring from one and two queens, respectively, suggested strongly that new queens do not produce a sexual brood until all workers of the old queen have died, which is consistent with kin selection theory.     

Amplitude of ecological potential: chub Leuciscus cephalus (L.) spawning in an artificial lowland canal

Chub Leuciscus cephalus (L.) is a rheophilic cyprinid which prefers lotic habitat conditions, shallow water (0.1–0.3 m) and gravel banks with moderate to high water flow (0.15–0.75m s^–1) for spawning. In contrast to these spawning requirements, a self-sustaining chub population was detected in a German lowland canal, the Oder–Havel Kanal. This canal is monotonously embanked with rip rap and almost completely lacks water flow and habitats that are suitable for riverine species. In 1999 chub spawned in the absence of water flow, in depths up to 1.28 m on allochthonous gravel with a mean diameter of 39 ± 16 mm (SD). These findings indicate a substantially higher environmental plasticity and tolerance of chub than previously reported.