The effect of neighbours on the mortality of harvester ant colonies

1. This study investigates how the density and size of conspecific neighbours affected the mortality of established colonies. Data were from a population of 250–300 colonies of the red harvester ant [ Pogonomyrmex barbatus (F. Smith)], on a 10-ha site, from 1988 to 1995.
2. Colonies are more likely to die as they grow older.
3. Small neighbours tend to lower the probability of dying.
4. The probability of dying varied significantly from year to year.
5. There was little effect of crowding on mortality.
6. There appeared to be no spatial effects, on the scale of 25. m, on the probability of dying.
7. Previous work shows effects of density on the founding of new colonies and on the reproductive output of established ones. The results presented here indicate that competition with neighbours rarely causes the death of established colonies.     

Queen lifespan and colony longevity in the ant Harpegnathos saltator

Abstract.  1. The longevity of field colonies was investigated in the ponerine ant Harpegnathos saltator (Jerdon) in which either reproductive workers (gamergates) or a single queen reproduce.
2. Data from 3 years were used to calculate the ratio between queen-right ( n  = 50) and gamergate ( n  = 12) colonies that can be used to derive the colony mortality of gamergate colonies. Using the survival rates of queens in the laboratory, extrinsic and intrinsic mortality rates of queen-right colonies were calculated.
3. No significant differences in the sizes of queen-right and gamergate colonies above 14 workers was found, suggesting that mortality of established colonies is not size related.
4. The mortality of gamergate colonies is 4.17 times higher than the intrinsic mortality of queen-right colonies.
5. In the laboratory, mean survival time of queens in colonies of more than 14 workers was 1.79 years.
6. Estimated mean survival time of queen-right and gamergate colonies in the field varies between 0.78 and 0.43 years respectively, when no costs of conflict during the replacement of queens occur; however, when the latter costs increase colony mortality to a level similar to extrinsic mortality, the calculated longevity of queen-right colonies would increase to 1.02 years.     

Serial monodomy in ants: an antipredator strategy?

Abstract.  1. The term serial monodomy is used to describe a life-history phenomenon in social insects. Serially monodomous colonies maintain multiple nests for their exclusive use, but only occupy one nest at a time.
2. The hypothesis that colony odours mediate nest relocation decisions was tested in the serially monodomous species Aphaenogaster araneoides from Central America. Odour extracts of colony members were created using a non-polar solvent.
3. Colonies strongly avoided reoccupying nests treated with colony odour extracts, while control colonies often returned to nests subjected to solvent-only control. Behavioural observations indicated that A. araneoides colonies are capable of detecting army ant ( Eciton burchellii ) raids up to 1.4 m from the raiding front, with several seconds to evacuate nests.
4. It is proposed that the function of serial monodomy in A. araneoides is the reduction of nest odour to enhance detection of predaceous army ants. Serial monodomy may be a widespread but undocumented mode of nesting where army ants occur in tropical and subtropical climates.     

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 .     

Temporal and spatial patterns in the emigrations of the army ant Dorylus (Anomma) molestus in the montane forest of Mt Kenya

Abstract.  1. The emigration behaviour of the army ant Dorylus ( Anomma ) molestus was studied in the montane forest of Mt Kenya. This species forages by massive swarm raids (mean width 10.3 m ± 4.6 m SD), which are assumed to have a strong negative impact on the densities of prey populations.
2. For non-reproductive colonies the stay duration in a nest is highly variable (median 17, range 3–111 days). This suggests that the frequency of emigrations is not dictated by a brood cycle as an underlying endogenous pattern generator.
3. Colony density is high (mean nearest neighbour's distance 82.1 m ± 29.4 m SD) and mean foraging range is 75.0 m, so encounters with neighbouring colonies occur frequently.
4. The straight line emigration distance is on average 92.7 m (± 29.7 SD). The emigration direction is random with respect to absolute bearing and also relative to the direction of the previous emigration. However, the emigration direction is influenced by the location of the nearest neighbour. Colonies typically emigrate directly away from their nearest neighbour.
5. Local food depletion is likely to be the ultimate cause for emigrations in this species, because emigration distance is larger than foraging range and colonies move away from their nearest neighbour. A small percentage of emigrations may be triggered by pangolin attacks on nests.
6. Contrary to the prediction of a recently developed mathematical model for epigaeic swarm-raiding Dorylus ( Anomma ) species, D.  ( A .)  molestus colonies do not engage in intraspecific battles. Possible reasons for the absence of fights between colonies despite apparently fierce intraspecific competition are discussed.     

The role of food and colony size in sexual offspring production in a social insect: an experiment

Abstract.  1. Large colonies of ants are known to have a higher propensity for sexual offspring production, probably owing to their high capacity to exploit food resources.
2. The effects of food supplementation on the propensity for sexual offspring production, and whether it is linked with colony size, were investigated in an environment with poor resources (clear-cut areas).
3. Large colony size was associated with a higher propensity for sexual offspring production in food-supplemented colonies, whereas in non-supplemented control colonies an association with colony size was not found.
4. The results demonstrate that large colonies seem to have a higher capacity to exploit supplemented food. In addition, the production of sexual offspring was apparently limited by food availability in clear-cuts, especially for large colonies.     

Effects of resource availability and social parasite invasion on field colonies of Bombus terrestris

Abstract.  1. The survival, growth and fecundity of bumblebee colonies are affected by the availability of food resources and presence of natural enemies. Social parasites (cuckoo bumblebees and other bumblebees) can invade colonies and reduce or halt successful reproduction; however, little is known about the frequency of invasion or what environmental factors determine their success in the field.
2. We used 48 experimental colonies of the bumblebee Bombus terrestris , and manipulated both resource availability at the landscape scale and date of colony founding, to explore invasion rates of social parasites and their effect on the performance of host colonies.
3. Proximity to abundant forage resources (fields of flowering oilseed rape) and early colony founding significantly increased the probability of parasite invasion and thus offset the potential positive effects of these factors on bumblebee colony performance.
4. The study concludes that optimal colony location may be among intermediate levels of resources and supports schemes designed to increase the heterogeneity of forage resources for bumblebees across agricultural landscapes.     

Habitat-specific life-history variation in the Caribbean termite Nasutitermes acajutlae (Isoptera: Termitidae)

Abstract.  1. Population and individual colony characteristics of Nasutitermes acajutlae on St John, U.S. Virgin Islands were studied over a 5-year period.
2. Four habitat types (dry forests, moist forests, sparse vegetation, and woodlands) were sampled for N. acajutlae , using quadrat-based methods. Lying dead wood was also quantified in the same sites.
3. Habitats differed significantly in the volume of lying dead wood, and total termite nest volume within quadrats was positively related to wood volume. It was hypothesised that the volume of dead wood affected growth, reproduction, and yearly survival of individual colonies.
4. In a sample of > 100 colonies assessed in 1998, 2000, 2001, and 2002, no significant differences in the growth rate of colonies (= increase in nest volume) could be found between habitat types; however, colonies in sparse vegetation were smaller, less likely to survive, and reproduced at a smaller size compared with those in the other three habitats.
5. Overall nest volume in sparse vegetation was high, compared with dry and moist forest, but colonies showed, on average, poorer survival (interpreted as lower fitness), and produced alates at an early stage of development. Optimum habitat characteristics for N. acajutlae may include not only favourable microclimate but also the reliable acquisition of new dead wood after storms.     

Colonization, riparian habitat selection and home range size in a reintroduced population of European beavers in the Loire

1. Colonization in a reintroduced population of European beavers in the Loire Valley was studied between 1974 and 1999. It followed a discontinuous remoteness model and a scattered distribution, beavers occupying only 25% of the river system over the 2800 km explored.
2. After 5 years, the colonization rate reached 104.2% year^–1 of new sites occupied (SD 75% year^–1), before dropping over the next 20 years. Nevertheless, the number of new colonies per km (0.125) remained stable throughout the years.
3.  Populus nigra , Salix alba and Fraxinus angustifolia were the dominant woody species in beaver sites, often associated with some herbaceous species.
4. The length of willow grove dominated by S. alba and P. nigra ( x ) was the best predictor of beaver home range ( y ), fitting the equation y =−0.742 x  + 5.9. Long-term maintenance of the population requires a minimum of 1.79-km of willows per colony.
5. In cutting tree trunks, beavers stimulate shoot development from the remaining stumps. They rejuvenate riparian forests, increase the number of tree stems and help stabilize the banks. Their effect on woody plant morphogenesis may have consequences for the helophyte communities used as food or habitat by other aquatic species.     

Colony productivity and foundress behaviour of a native wasp versus an invasive social wasp

Abstract.  1. Colony productivity, prey utilisation, and foundress behaviour of a North American native wasp ( Polistes fuscatus ) versus an European invasive wasp ( Polistes dominulus ) were investigated in a controlled field experiment with optimal versus natural foraging conditions. Colonies with the optimal prey foraging conditions were provided with prey ad libitum within an enclosed area. The other colonies foraged in the adjacent field–woodland but had the same nest conditions as the other treatment.
2. When given prey ad libitum , both wasp species captured similar amounts of prey and the conversion to total offspring biomass was similar. But P. dominulus colonies produced 2.5 times the number of workers as P. fuscatus colonies, reflecting the smaller size of P. dominulus wasps.
3. Foundresses of P. dominulus were observed more often building or repairing the nest, thereby contributing to the production of colonies with twice as many cells as colonies of P. fuscatus . Foundresses of P. dominulus showed more acts of aggression toward workers than did P. fuscatus foundresses, which was not a function of adult density on the nest.
4. At the end of the experiment, P. dominulus colonies with optimal prey foraging conditions still had a high level of egg-laying and peaked in the number of pupae then, whereas egg-laying and the number of pupae per colony of the other treatments began to decline 2–3 weeks earlier. These results indicate that P. dominulus is more opportunistic than P. fuscatus , which may account in part for P. dominulus 's success as an introduced species in North America.     

Dynamic matching of forager size to resources in the continuously polymorphic leaf-cutter ant, Atta colombica (Hymenoptera, Formicidae)

Abstract.  1. Ergonomic optimisation theory proposes that by increasing variation in worker morphology, social insect colonies may increase their dietary breadth; however, little is known about how this relationship operates at the colony level. This study examines the colony-level pattern of forager size allocation to resource sites in a natural setting.
2. Using a biologically relevant measure of toughness, it is shown that leaf-cutter ant colonies exploit a variety of plant resources that vary significantly in toughness at any given time.
3. Forager size is shown to be matched to the toughness of plant material, with larger ants harvesting tougher material.
4. Furthermore, outbound foragers travelling to a harvest site are matched in size to the toughness of plant material contained within the site and are not a random selection of available foragers. The match between forager size and plant toughness may reduce the number of wasted trips and ill-matched foragers.
5. The observed colony-level pattern of forager allocation could be the result of learning by individual foragers, or the result of information shared at the colony level.     

Recruitment in the harvester ant, Pogonomyrmex occidentalis: effects of experimental removal

We examined the importance of experimental removal of mature colonies on colony recruitment in the western harvester ant Pogonomyrmex occidentalis. To test the common assumption that established colonies suppress the establishment of new colonies we removed all colonies in ten 0.25 ha plots in 1996 and an additional five plots in 1997 and measured new colony recruitment in 1997, 1998, and 1999. Using a blocked, paired plot design we found that removal of colonies increased new colony recruitment in some areas of the site, but not others. Spatial variation in the importance of established colonies to recruitment was consistent across years; blocks in which density dependence was important in one year exhibited density dependent recruitment in following years. We estimated that in the blocks where recruitment was affected by established colonies, they accounted for less than 10% of the mortality of foundress queens. The increase in the number of new recruits (on average two additional new colonies) was considerably less than the number of colonies removed; average colony density in the removal plots was 14 colonies per 0.25 ha plot. The consistent lack of importance of established colonies to recruitment in one block and the relatively small response to colony removal in the other blocks suggests that the number of new colonies in a year may not be equivalent to the number of deaths of established colonies in that year. Space limitation is an important influence on recruitment in P. occidentalis, but the magnitude of the limitation varies spatially.     

Advantages of multiple foundress colonies in Belonogaster juncea juncea L.: greater survival and increased productivity

Abstract.  1. The ecological hypothesis predicts that multiple foundress colonies of social wasps may have a better survival rate and produce more brood per capita than single foundress colonies. With the aim of verifying if these characteristics exist in the primitively eusocial wasp species Belonogaster juncea juncea (L.), we monitored 49 foundations, including 13 single and 36 multiple foundress colonies, in Cameroon.
2. Multiple foundress colonies were significantly more successful than single foundress colonies in producing at least one adult.
3. The total productivity of the colonies increased significantly with the number of associated foundresses, but the productivity per capita did not. No single foundress colony reached the sexual phase, while eight (21.6%) multiple foundress colonies did. Males were produced in only five colonies, so that the sex ratio was biased in favour of females.
4. These results suggest that because of the strong ecological constraints on solitary nesting, survival and high colony productivity are two advantages of multiple foundress colonies in B. j. juncea .
5. The decreasing per capita productivity concomitant with an increasing number of females noted in this study illustrates once again Michener's paradox. The coefficient of variance of the per capita productivity significantly decreased with group size, as Wenzel and Pickering suggested in the model they created to explain the paradox.
6. Ecological factors may act in conjunction with other factors, such as genetic relatedness between associated foundresses, to promote joining behaviour in B. j. juncea .     

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.     

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.     

Relationships between Long-Term Demography and Weather in a Sub-Arctic Population of Common Eider

Effects of local weather on individuals and populations are key drivers of wildlife responses to climatic changes. However, studies often do not last long enough to identify weather conditions that influence demographic processes, or to capture rare but extreme weather events at appropriate scales. In Iceland, farmers collect nest down of wild common eider Somateria mollissima and many farmers count nests within colonies annually, which reflects annual variation in the number of breeding females. We collated these data for 17 colonies. Synchrony in breeding numbers was generally low between colonies. We evaluated 1) demographic relationships with weather in nesting colonies of common eider across Iceland during 1900–2007; and 2) impacts of episodic weather events (aberrantly cold seasons or years) on subsequent breeding numbers. Except for episodic events, breeding numbers within a colony generally had no relationship to local weather conditions in the preceding year. However, common eider are sexually mature at 2–3 years of age and we found a 3-year time lag between summer weather and breeding numbers for three colonies, indicating a positive effect of higher pressure, drier summers for one colony, and a negative effect of warmer, calmer summers for two colonies. These findings may represent weather effects on duckling production and subsequent recruitment. Weather effects were mostly limited to a few aberrant years causing reductions in breeding numbers, i.e. declines in several colonies followed severe winters (1918) and some years with high NAO (1992, 1995). In terms of life history, adult survival generally is high and stable and probably only markedly affected by inclement weather or aberrantly bad years. Conversely, breeding propensity of adults and duckling production probably do respond more to annual weather variations; i.e. unfavorable winter conditions for adults increase probability of death or skipped breeding, whereas favorable summers can promote boom years for recruitment.     

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.     

Spread of plague among black-tailed prairie dogs is associated with colony spatial characteristics

Sylvatic plague (Yersinia pestis) is an exotic pathogen that is highly virulent in black-tailed prairie dogs (Cynomys ludovicianus) and causes widespread colony losses and individual mortality rates >95%. We investigated colony spatial characteristics that may influence inter-colony transmission of plague at 3 prairie dog colony complexes in the Great Plains. The 4 spatial characteristics we considered include: colony size, Euclidean distance to nearest neighboring colony, colony proximity index, and distance to nearest drainage (dispersal) corridor. We used multi-state mark–recapture models to determine the relationship between these colony characteristics and probability of plague transmission among prairie dog colonies. Annual mapping of colonies and mark–recapture analyses of disease dynamics in natural colonies led to 4 main results: 1) plague outbreaks exhibited high spatial and temporal variation, 2) the site of initiation of epizootic plague may have substantially influenced the subsequent inter-colony spread of plague, 3) the long-term effect of plague on individual colonies differed among sites because of how individuals and colonies were distributed, and 4) colony spatial characteristics were related to the probability of infection at all sites although the relative importance and direction of relationships varied among sites. Our findings suggest that conventional prairie dog conservation management strategies, including promoting large, highly connected colonies, may need to be altered in the presence of plague. © 2011 The Wildlife Society     

Nourishment affects colony demographics in the paper wasp Polistes metricus

Abstract  1. Colony survivorship and numbers of nest cells, pupae, and adult females were monitored throughout the nesting season for a cohort of 78 colonies of the paper wasp Polistes metricus Say. Thirty-nine colonies received a twice-weekly nourishment supplement of honey during pre-emergence and early emergence phases of the colony cycle; 39 colonies were unsupplemented controls.
2. Colony survivorship was unaffected by the supplemental nourishment. Loss of colonies to predation differed among three sites but was unaffected by supplementation.
3. Honey-supplemented colonies constructed more nest cells than did control colonies but this effect was not expressed until after supplementation had ceased.
4. Honey-supplemented colonies produced more pupae than did control colonies but the number of adult females at nests did not differ between supplemented and control colonies. Because honey-supplemented colonies had more offspring but fewer of them remained as workers at the nest, honey supplementation led to a lower frequency of workers and corresponding higher frequency of reproductives than in control colonies.
5. In a second year of study, colony survivorship and numbers of nest cells, pupae, and adult females were monitored from late pre-emergence until the end of the nesting season for a cohort of 32 colonies of Polistes metricus . In 16 colonies, trophallactic saliva was taken from final-instar larvae on nine dates in the late pre-emergence and early emergence periods; 16 colonies served as controls.
6. Saliva-diminished colonies had lower survivorship, fewer nest cells, fewer pupae, and fewer adult females at the nest than did control colonies.
7. These results show that variation in nourishment in the early to mid phases of the colony cycle can have significant effects on the subsequent colony demographics of Polistes metricus paper wasps.     

Delayed maturation in the colonial coralGoniastrea aspera (Scleractinia): whole-colony mortality, colony growth and polyp egg production

The present study examines (1) the cost of reproduction on colony growth, and (2) relationships among sexual maturity, whole-colony mortality rate and colony growth rate inGoniastrea aspera free from external influences by macrobenthos. Survival of colonies in permanent plots was followed for two years. Egg production by polyps in colonies collected just before the first spawning of a year was estimated by dissecting the polyps. Growth of the colonies (increase in number of polyps) was followed over one annual reproductive cycle. The cost of egg production on colony growth was apparent through colony ontogeny: (1) immature colonies had a greater annual growth rate than mature colonies, but produced almost no eggs; (2) in mature colonies, growth rate was negatively correlated with NE/PV (number of eggs per polyp volume mm^-3). Annual whole-colony mortality was high in colonies with fewer than11 polyps in initial colony size, while mortality was extremely low once a colony grew beyond this size. This critical size for low whole-colony mortality was much smaller than the colony size (40 polyps) which would attain maturity one year later. Age at maturity was estimated as six years. While survival to maturity may be a selective force for the evolution of delayed maturation, the present data suggest that high colony fecundity, achieved after a long growth period as an immature colony, and an abrupt decrease of colony growth rate after maturation are the crucial forces.