Phenotypic variation in male and worker encapsulation response in the bumblebee Bombus terrestris

Abstract.  1. Life-history traits such as immunity are often characterised by the presence of large phenotypic variation, but it often remains unclear how and why this variation is maintained by selection.
2. Here an annual social insect, the bumblebee Bombus terrestris , was used to study variation in encapsulation response of males and workers. Bumblebees are a suitable system to study offspring immunity because they are host to a broad variety of different parasites. Bumblebee males, in particular, have a long lifespan compared with other social insect males and their immunity should therefore be an important element for colony reproductive success.
3. Encapsulation response, which was used here as a measurement for the generalised immune defence capacity of an individual, was found to be a highly variable trait. High levels of worker response correlated with low levels of colony parasitism rates.
4. Encapsulation response was found to be (a) lower in males compared with sister workers, and (b) lower in late-produced cohorts compared with early ones.
5. In colonies with delayed sexual reproduction, males had a lower encapsulation response. Thus, investments into immunity seemed reduced in later male cohorts and those eclosing later in the season, perhaps because males had a shorter expected remaining time to acquire matings. The results presented add further evidence that immune defence is a key variable defining colony fitness in social insects.     

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.     

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.     

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 .     

Effects of reproductive timing and colony size on the survival, offspring colony size and drone production in the honey bee (Apis mellifera)

ABSTRACT. 1. The effects of colony size and time of reproduction on the survival and size of offspring colonies and on drone production were examined for honey bees, Apis mellifera L. Drone and worker production and survival of parental and offspring colonies were monitored following swarming. Also, the temporal patterns of drone emergence and availability of unmated queens were examined.
2. Colony size at swarming was positively correlated with the number of workers invested in offspring colonies and the number of queens produced. However, colony size at swarming was not correlated with the number of offspring colonies produced.
3. Swarm size was positively correlated with drone and worker production after swarms were hived. Worker production of hived swarms was positively correlated with colony survival. Offspring queens which inherited a parental nest survived longer than queens in either primary swarms or afterswarms, presumably due to the advantage of inheriting a nest.
4. Drone emergence peaked just prior to swarming, the time when unmated queens were available. High drone production by colonies initiated by swarms probably reflected an attempt to reproduce prior to winter. The probabilities of a second swarming cycle within the same year and of surviving the winter were low for colonies initiated from swarms.     

Resource characteristics and competition affect colony and individual foraging strategies of the wood ant Formica integroides

Abstract.  1. Resource characteristics and competitive pressure can affect an ant colony's foraging strategy. This study examined the ability of the wood ant Formica integroides to respond, at both the colony and individual levels, to changes in competitive pressure for access to terrestrial and arboreal resources.
2. Because foraging behaviours depend on resource characteristics, foraging for different resource types (e.g. terrestrial and arboreal habitats) produces different spatial or territorial arrangements. In this study, terrestrial contests for resources followed an interference-exploitation tradeoff, while arboreal foragers defended entire trees as absolute territories.
3. Competitive pressure for access to arboreal resources was shown to increase with distance from F. integroides nests.
4. In this study, the ability of F. integroides to defend a resource varied with body size. Large foragers were better defenders than small foragers. For groups of foragers, the ability to defend a resource increased with the ratio of large to small foragers.
5. In response to competitive pressure, F. integroides colonies altered the size distribution of arboreal, but not terrestrial, foragers. An increase in competitive pressure was matched by an increase in the number of large foragers allocated to trees. This response to competition affected the relationship between body size and distance from the nest for arboreal foragers.
6. Foraging behaviours for individual arboreal foragers also varied with competitive pressure. As competition increased, large arboreal foragers spent more time in direct contact with the resource rather than standing between resource patches.     

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.     

Colony pollen reserves affect body size,sperm production and sexual development in males of the stingless bee <Emphasis Type="Italic">Melipona beecheii</Emphasis>

The production of male sexual offspring by social insect colonies is often strongly seasonal or resource-dependent. In stingless bees, males are produced in smaller numbers under conditions of low colony food reserves; whether such males are negatively affected in traits related to reproductive success is not known. We compared body size, sperm production and sexual maturity in Melipona beecheii males reared with experimentally supplemented or reduced pollen reserves, but with otherwise equal numbers of workers and equal quantities of honey reserves. We also studied the same traits in males collected from non-manipulated colonies with pollen reserves intermediate between the supplemented or reduced groups but with more workers and honey reserves. Males reared under experimentally reduced pollen reserves had significantly smaller bodies and lower sperm counts compared to those reared in colonies with experimentally supplemented pollen reserves. There was also a significantly positive relationship between the number of sperm and body size in males across all colony treatments. The maximum number of sperm in seminal vesicles was recorded 2 days later in males from colonies with reduced pollen compared to males from colonies with supplementary pollen. Males from non-manipulated colonies were intermediate in size, sperm count and speed of maturation. Our study documents for the first time the existence of large size variation in males of stingless bees that is related with the amount of pollen reserves in their natal colony. We conclude that a colony’s pollen reserves have a major impact on male body size, sperm production and speed of sexual maturity in this stingless bee, which may be the case in other social insects. Stingless bees are a good model system to study the balance between colony-level selection and individual-level selection on male sexually selected traits such as body size.     

Reproductive strategy of the slave antFormica podzolica relative to raiding efficiency of enslaver species

Summary Formica podzolica serves as host to slave-making ants in North America. We propose thatF. podzolica may respond to slavery by two alternative colony-growth and reproductive strategies depending on the raiding ability of the slavemaker: (1) Rapid colony growth at the expense of producing sexuals to a stage where raiding by unspecialized, facultative slavemakers, capable of exploiting only small colonies, becomes unlikely owing to a strong work force and (2) Early production of sexual offspring at the cost of colony growth to secure some sexual production in an environment with specialized obligate enslavers, capable of raiding large colonies. We tested the strategies by excavating 30 small to moderately large mounds ofF. podzolica and measured reproductive parameters of colonies in relation to mound size, worker number, and worker size. Mound area predicted worker number satisfactorily. Worker number correlated significantly with worker head width and with number of worker and sexual offspring. With a growing work force, the proportion of sexual offspring increased in the total offspring. Two thirds of the colonies producing sexuals emitted single sex, sex being independent of colony size. Some of the large colonies produced both sexes with a strong bias toward either sex. The unweighted population-level sex ratio did not differ from even, being 0.52 (numerical) or 0.54 (biomass). Very large mounds (not excavated) had small workers and highly male-biased sex ratios, probably owing to energy constraints set by central-place foraging. Population-level colony ontogeny data did not fit either one of the suggested strategies, but imply a mixture of the two. We discuss an alternative, still untested raid-independent explanation to the ontogeny pattern.     

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 .     

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.     

Sexual size dimorphism in the two spot ladybird beetle Adalia bipunctata: developmental mechanism and its consequences for mating

Abstract  1. The literature on ladybirds indicates that males are consistently smaller than females but take the same length of time to complete their development. Rearing Adalia bipunctata at 20 and 25 °C confirmed that protandry cannot account for sexual size dimorphism in this species, nor can a difference in egg size.
2. Female larvae consumed more food and had a higher relative growth rate in the fourth instar than did male larvae.
3. When food is limited, small males appear to be more successful at mating than are large males.
4. To account for these results, it is hypothesised that the gonads of male larvae compete more strongly with the soma for resources and that this reduces the growth potential of the soma of male larvae relative to that of female larvae. The greater mating success of small males when food is limited supports the eat or mate hypothesis, which predicts that when food is limited small males will spend less time feeding and more time mating than will large males.     

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.     

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.     

Three clonal types of urothelium with different capacities for replication

Objectives:  Similar to other epithelia, urothelium in vivo has a hierarchal organization of cells each with specific gradients of differentiation. While distinct cell types have been described as important in bladder cancer in vitro , clonal and proliferative capacities of normal urothelial cells have not been characterized.
Materials and methods:  Three cell types and colony types were identified from primary porcine urothelial culture. Proliferative activity, patterns of apoptosis and differentiation, colony forming efficiency and ability to change phenotype with passage were determined and compared.
Results:  Small, T-I colonies with large flattened (type-1) cells had low levels of proliferation and high levels of apoptosis. Large T-III colonies had a central area of small (type-3) cells surrounded by type-1 and type-2 cells. Proliferation and apoptosis were asymmetrically distributed in the periphery of T-II and T-III colonies. T-III colonies proved to be significantly more clonogenic and proliferative. With appropriate induction, type-1 cells were able to proliferate upon passage and form type-3 cells, yet long-term culture demonstrated that progeny of type-1 cells appeared to have inherited a clonogenic handicap.
Conclusions:  Type-3 cells in the centre of T-III colonies appear to harbour stem-like qualities with a relatively low proliferative and apoptotic index at homeostasis and the ability to become highly proliferative upon passage. This study demonstrates that distinct urothelial cell types with differing clonal capacities can be isolated from the bladder and these cells may have implications for tissue engineering and carcinogenesis.     

Spatial and temporal variation in a suite of life-history traits in two species of wolf spider

Abstract.  1. Life-history traits and density were assayed in seven populations of two sympatric species of wolf spider for three consecutive years. The goal of the study was to quantify population dynamics and its relation to spatial and temporal life-history variation.
2. Adult female body size and fecundity varied significantly, among field sites and among years, in both species. Female spiders of both species differed in mean relative reproductive effort among sites, but not among years. The size of offspring was invariable, with no significant differences due to site or year.
3. All populations of both species tended to either decrease or increase in density during a given year and this was tightly correlated with changes in prey consumption rates.
4. Since life-history patterns are determined primarily by selection, it is concluded that size at sexual maturity for females is phenotypically plastic and responds to changes in prey availability. Offspring size however is not plastic and it is likely that other selection forces have determined offspring size. Temporal fluctuations in population size are correlated over a large area relative to dispersal capabilities for these species and conservation efforts for invertebrates must take this into consideration.     

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.     

Influence of adult nutrition on the relationship between body size and reproductive parameters in a parasitoid wasp

Abstract.  1. An important constraint upon life-history evolution in parasitoids is the limit imposed by body size on allocation of limited metabolic resources to different fitness-related physiological functions such as reproduction and survival.
2. The influence of adult nutrition on reproductive and maintenance variables was studied in the synovigenic ectoparasitoid Mastrus ridibundus , and it was determined whether resource allocation to these different functions depends on body size.
3. Over the course of adult life there was a positive relationship between body size and the number of mature eggs in adult females both in the presence and absence of food. However, only in the presence of food did egg maturation rates increase significantly with body size. Starved wasps produced significantly smaller eggs than fed ones, which has not been documented before. Moreover, starved wasps produced fewer offspring than fed wasps, and attacked fewer hosts.
4. The availability of food had a major effect on longevity, with fed females living about 10 times longer than starved ones. There was also a positive relationship between body size and longevity. In starved wasps, this relationship was the same both in the presence and absence of hosts, but in fed wasps there was a positive relationship between body size and longevity in the absence of hosts only. Allocation to initial eggs relative to lifetime progeny production did not decline with body size.
5. The data reveal that in M. ridibundus the trade-off between maintenance and reproduction varies with life expectancy.     

Alarm pheromone emission by pea aphid, Acyrthosiphon pisum, clones under predation by lacewing larvae

Genetic variation in anti-predator traits has been shown for a variety of species. Aphid alarm pheromone, ( E )-β-farnesene, is released by attacked aphids and causes a variety of behavioral defense reactions in the signal receivers. In pea aphids, Acyrthosiphon pisum Harris (Homoptera: Aphididae), ( E )-β-farnesene mediates the production of winged offspring in the presence of natural enemies. While variation in the propensity for pea aphids to produce winged offspring is well-documented, little quantitative information is available about clonal differences in ( E )-β-farnesene emission or the amount of alarm pheromone released in aphid colonies. We tested the wing induction response of four clones when attacked by a predatory lacewing larva, Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae), and found that three of the four clones increased the proportion of winged offspring under predator attack. We then investigated the emission of aphid alarm pheromone of these clones of pea aphid under attack. Alarm pheromone emission in aphid colonies of initially 25 adults varied from 81.2 to 10 851.0 ng per aphid colony over 24 h. There were no differences between clones in total emission or in emission per consumption event. These results show that there is substantial variability in alarm pheromone emission within clones and that the propensity to produce winged offspring in some clones is not a simple function of the propensity of alarm pheromone production in these clones.     

On foraging,recruitment systems and optimum number of scouts in eusocial colonies

Summary A numerical model of an eusocial colony foraging for food showed that, for each set of values of resource density, resource size and recruitment system employed, a given optimal proportion of scouts in the colony maximize the amount of resources retrieved by a colony during a fixed period. The model predicts that ants using mass recruitment systems should have larger colonies with small foragers, and should forage on large food sources. Retrieval of small food sources by small colonies is best achieved with large workers using individual foraging strategies. For mass foragers, several food sources are best retrieved using democratic decision-making systems in recruitment, whereas for very large food sources at very low mean food patch density, autocratic decision-making systems are optimal. Some of the experimental evidence available is discussed in the light of these findings, as they confirm the prediction that large colonies with small workers have mass recruitment systems, whereas workers of small colonies with large workers are generally lone foragers.