Reproductive schedule and factors affecting soldier production in the eusocial bamboo aphid Pseudoregma bambucicola (Homoptera, Aphididae)

Summary: The reproductive characteristics of the soldier-producing aphid Pseudoregma bambucicola were studied in Kagoshima, Southern Japan, to know the factors affecting soldier production of eusocial aphids. The soldier proportion in aphid colonies was highest from October to November. In some large colonies, soldiers were observed in all seasons except in July when colony size was relatively small. Multiple regression analysis showed that the colony size was a principal factor affecting soldier proportion throughout a year. Other social or environmental factors such as aphid composition, host plant conditions and predator abundance were not always significant. Rearing experiments revealed that large colonies (̿,000 individuals) produced soldiers in almost all seasons while small colonies (<1,000) never produced any soldiers. The caste-production schedule of adult females was examined in the field. When solitary females produced both castes, they usually produced normal nymphs first and then soldiers. Females from large colonies tended to produce more soldiers in the earlier period of their lifetime, whereas females from newly established small colonies produced no or only a few soldiers at later times. The average number of soldiers and normal nymphs produced consecutively by a single female was >10 and >20, respectively. Because they have a small number of ovarioles (<15 on average), females should alter caste production within the same ovarioles according to changes in environmental conditions. Artificial removal or introduction of predators and reduction of colony size did not affect soldier production over two successive generations, revealing maternal effects on soldier production. Females cannot shift caste production quickly in response to changes in predator abundance and colony size. This is probably due to early developmental determination of castes within the mother's body.     

Correlates of reproductive success among field colonies of Bombus lucorum: the importance of growth and parasites

Abstract.

• 1 In natural populations, colonies of bumble bees vary in many important life history traits, such as colony size and age at maturity, or the number and sex of reproductives produced. We investigated how the presence of parasites in field populations of the bumble bee Bombus lucorum L. relates to variation in life history traits and reproductive performance. A total of thirty-six colonies was placed in accessible nest sites in the field and monitored at regular intervals throughout a season.
• 2 Among the life history correlates, early nest foundation was strongly associated with large maximum colony size, old age and large size at maturity, and this in turn with successful production of males and queens, as well as with the number of sexuals produced. Overall, reproductive success was highly skewed with only five colonies producing all the queens. Sixteen colonies failed to reproduce altogether.
• 3 The social parasite Psithyrus was abundant early in the Bombus colony cycle and preferentially invaded host nests with many first brood workers and thus disproportionately large size, i.e. those colonies that would otherwise be more likely to reproduce or produce (daughter) queens rather than males. To prevent nest loss, Psithyrus had to be removed soon after invasion. Therefore, the effects reported here can only be crude estimates.
• 4 Parasitoid conopid flies are likely to cause heavy worker mortality when sexuals are reared by the colonies. Their inferred effect was a reduction in biomass that could be invested in sexuals as well as a shift in the sex ratio at the population level resulting from failure to produce queens. We suggest to group the inferred correlates into ‘early events’ surrounding colony initiation and social parasitism, and ‘late events’ surrounding attained colony size in summer and parasitism by conopid flies. Our evidence thus provides a heuristic approach to understand the factors that affect reproductive success of Bombus colonies.

     

Mating strategy and isolation between the two forms, macrogyna and microgyna, of Myrmica ruginodis (Hym. Formicidae)

Abstract.

• 1 It is shown that the size distribution and degree of overlap between individual queens of the two forms of Myrmica ruginodis, microgyna and macrogyna, is exactly the same in southern England as was originally described from Scotland.
• 2 Distinguishing colonies of the two forms is not as easy as distinguishing between individuals, both types can be polygynous and approximately 25% of colonies contain a mixture of queens.
• 3 Males of the two forms can be distinguished solely on the basis of size and, in general, macrogyna colonies produce larger males, workers and gynes than microgyna colonies. There is little correlation between the sizes of males, workers and gynes in colonies within the macrogyna and microgyna groups.
• 4 If the two forms are separate species and mixing is a parasitic association (as suggested elsewhere), then there should be breeding isolation between the forms. This is tested by examining data from nine mating-swarms.
• 5 All the swarms contain a significant proportion of the microgyna form. There is no evidence of assortative mating, although larger males are more likely to get a mate than smaller ones. This behaviour, combined with the possibility that microgynes mate near to the nest, might prevent complete mixing during mating.
• 6 The status of the forms is discussed. Besides the possibility of a‘pre-parasitic’relationship it is suggested that the forms might represent a polymorphism, present in all populations, the balance between them being the result of selection determined by local environmental factors.

     

Fire ant predation on native and introduced subterranean termites in the laboratory: effect of high soldier number in Coptotermes formosanus

Abstract.

• 1 Native Reticulitermes spp. Holmgren and the introduced Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) were exposed to predation in the laboratory by Solenopsis invicta Buren (Hymenoptera: Formicidae), a known predator of Reticulitermes.
• 2 In one experiment, both taxa contained the 2.4% soldiers commonly found in Reticulitermes. In a second experiment, Reticulitermes contained 2.4% and C.formosanus a more normal 18.3% soldiers.
• 3 When soldier proportions were equal, C.formosanus soldiers suffered less of a decrease in survivorship due to predation compared with Reticulitermes soldiers. Little difference was found between taxa in the effect of predation on termite workers or in the effect of termites on predator survivorship.
• 4 When soldier proportions were unequal, both C.formosanus castes suffered less from predation than those of Reticulitermes. Again, little difference was found in the effect of termites on ants.
• 5 C.formosanus showed less movement into new containers when soldier number was abnormally low, compared with both taxa with normal caste proportions.

     

Social structure and the defensive role of soldiers in a eusocial bamboo aphid,Pseudoregma bambucicola (Homoptera: Aphididae): A test of the defence-optimization hypothesis

A complete understanding of the evolution of sociality in aphids requires a detailed knowledge of the patterns of soldier investment in their ecology. The eusocial bamboo aphidPseudoregma bambucicola has a morphologically specialized first-instar soldier caste. The proportion of soldiers was positively correlated with colony size. Within a colony, soldiers were evenly distributed among subcolonies; within each subcolony, however, their distribution was biased toward peripheries which were exposed to many predators. Field experiments introducing natural enemies such asEupeodes confrater (Diptera: Syrphidae) andSynonycha grandis (Coleoptera: Coccinellidae) revealed that the survival rate of these predators was negatively correlated with the density of soldiers, suggesting that soldiers can more or less defend their colonies by killing or removing a range of natural enemies. Observations suggest that large mature colonies attract more predators than newly established small colonies and that, within a colony, the predators attack each subcolony regardless of its position on bamboo shoots. This implies the presence of a positive correlation between colony size and predation risk. Thus, the investment in soldiers seems to reflect the attacking pattern of predators within a colony. These results agree with the defence-optimization hypothesis in soldier investment ofP. bambucicola colonies.     

Keeping a low profile: small hive beetle reproduction in African honeybee colonies

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Successful and unsuccessful development of colonies of Vespula vulgaris (Linn.) (Hymenoptera: Vespidae)

Abstract.

• 1 Six categories of immature and five categories of mature colonies are described and their developmental relationships suggested.
• 2 Mature brood destroyed by the workers became numerous in 46.4% of the immature colonies. Most of these formed unsuccessful colonies (36.9%) producing few or no queens. The remaining 9.5% recovered to form successful colonies.
• 3 Successful colonies either reared queens only (mean 356 queens), or queens (mean 1461) and males (mean 254) in their large cells.
• 4 The number of small cells in mature colonies correlated with the number of cells built by the founder queen.
• 5 The greater the number of large cells in a mature colony the greater were the chances that a larger proportion of these cells would be used to rear adults.
• 6 The number of small and large cells are correlated in mature successful colonies.
• 7 Queen activity and quality are important factors in the control of colony size and success.
• 8 Selective destruction of male sealed brood in the large cells is linked to worker, rather than queen, control.

     

Size‐structured vulnerability of the colonial cyanobacterium,Microcystis aeruginosa,to grazing by zebra mussels (Dreissena polymorpha)

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Colony founding in the primitively eusocial wasp, Ropalidia marginata (Hymenoptera: Vespidae)

Abstract.

• 1 In a 16-month study in Bangalore, India, about 35% of the newly founded colonies of Ropalidia marginata were single foundress colonies and the remainder were multiple foundress colonies with two to twenty-two individuals.
• 2 Larger colonies did not have a significantly higher per capita productivity, did not produce significantly heavier progeny and did not produce them significantly faster than smaller colonies did.
• 3 Predation by the hornet Vespa tropica appeared to be independent of group size.
• 4 Single foundress colonies failed more often but not often enough to make them have a lower average per capita productivity, compared to multiple foundress colonies.
• 5 Some of the advantages of multiple foundress associations came from the greater predictability of their attaining the mean per capita productivity, the relatively lower rates of usurpation experienced by them compared to single foundress colonies, and the opportunities provided by queen turnovers for workers to become replacement queens and gain direct individual fitness.

     

Community structure and antifungal activity of actinobacteria in a fungus-growing termite

1. Fungus-growing termites cultivate the fungal mutualist Termitomyces as their main food source; however, how fungus-growing termites protect Termitomyces from threats is still unclear. In this study, we investigated the actinobacterial communities in Odontotermes formosanus individuals and their fungal combs.
2. Moreover, the antifungal activities of the isolated actinobacteria were tested. 16 S rRNA gene sequencing results indicated that the actinobacteria in O. formosanus and its fungal combs belong to 5 classes, 17 orders, 40 families, and 84 genera.
3. The relative abundance of Coriobacteriia in the nymphs, young workers, old workers, and soldiers was higher than that in the queens and fungal combs, and the relative abundance of class Actinobacteria in the queens and the fungal combs was higher than that in the nymphs, young workers, old workers, and soldiers.
4. Based on antifungal bioassays, 3 strains of Amycolatopsis and 2 strains of Streptomyces isolated from old workers had strong inhibitory activity against Xylaria angulosa but weak inhibitory activity against Termitomyces sp.
5. These results indicated that the actinobacteria of O. formosanus may contribute to protecting termite fungal food via their asymmetric antifungal activities.

     

Interaction between flight,reproductive development and oviposition in the pine weevil Hylobius abietis

• 1 The development of reproductive and flight capacity of pine weevils Hylobius abietis during the spring and their dispersal to, and subsequent development at, new clearfell oviposition sites comprise key phases in their life cycle in managed forests. At an old clearfell site where autumn‐emerging weevils had overwintered, weevils were trapped as they re‐emerged in the spring and tested for their ability to fly and then dissected to determine the degree of wing muscle and egg development.
• 2 Re‐emerging weevils were most abundant in pine growing at the edge of the clearfell and, over most of the trapping period (April to June), their capacity for flight (proportion flying and wing muscle width) was more advanced than in weevils from the clearfell itself, with a similar trend in the degree of reproductive development (proportion with mature eggs and egg volume).
• 3 In weevils from the clearfell, flight capacity and reproductive development increased concurrently to a peak around mid‐May. In weevils from pine, wing muscles were already well developed at the start of trapping, although few of them flew. Their more advanced development was attributed to the increased opportunities for maturation feeding after emergence in the previous autumn.
• 4 In the spring, weevils reached the canopy of trees for maturation feeding by walking and, to a lesser extent, by flight. Weevils dispersed by flight to oviposition sites in mid‐May when most of them were reproductively mature. After arrival, flight ability and wing muscle size declined rapidly but egg production was maintained until most weevils had stopped flying. When wing muscles reached their minimum size, there was a marked decline in egg size, suggesting that wing muscle breakdown is important in maintaining egg production at oviposition sites. Prospects for further wing muscle and reproductive development are discussed.

     

Tradeoffs in the Evolution of Caste and Body Size in the Hyperdiverse Ant Genus Pheidole

The efficient investment of resources is often the route to ecological success, and the adaptability of resource investment may play a critical role in promoting biodiversity. The ants of the “hyperdiverse” genus Pheidole produce two discrete sterile castes, soldiers and minor workers. Within Pheidole, there is tremendous interspecific variation in proportion of soldiers. The causes and correlates of caste ratio variation among species of Pheidole remain enigmatic. Here we test whether a body size threshold model accounts for interspecific variation in caste ratio in Pheidole, such that species with larger body sizes produce relatively fewer soldiers within their colonies. We evaluated the caste ratio of 26 species of Pheidole and found that the body size of workers accounts for interspecific variation in the production of soldiers as we predicted. Twelve species sampled from one forest in Costa Rica yielded the same relationship as found in previously published data from many localities. We conclude that production of soldiers in the most species-rich group of ants is regulated by a body size threshold mechanism, and that the great variation in body size and caste ratio in Pheidole plays a role in niche divergence in this rapidly evolving taxon.     

You'd better walk alone: Changes in forest composition affect pollination efficiency and pre‐dispersal cone damage in Iberian Juniperus thurifera forests

• Changes in land‐use patterns are a major driver of global environmental change. Cessation of traditional land‐use practices has led to forest expansion and shifts in forest composition. Consequently, former monospecific forests maintained by traditional management are progressing towards mixed forests. However, knowledge is scarce on how the presence of other tree species will affect reproduction of formerly dominant species. We explored this question in the wind‐pollinated tree Juniperus thurifera. We hypothesised that the presence of heterospecific trees would have a negative effect on cone production and on the proportion of cones attacked by specialised predators.
• We assessed the relative importance of forest composition on cone production, seed development and pre‐dispersal cone damage on nine paired pure and mixed J. thurifera forests in three regions across the Iberian Peninsula. The effects of forest composition on crop size, cone and seed characteristics, as well as damage by pre‐dispersal arthropods were tested using mixed models.
• Cone production was lower and seed abortion higher in mixed forests, suggesting higher pollination failure. In contrast, cone damage by arthropods was higher in pure forests, supporting the hypothesis that presence of non‐host plants reduces damage rates. However, the response of each arthropod to forest composition was species‐specific and the relative rates of cone damage varied depending on individual tree crops.
• Larger crop sizes in pure forests compensated for the higher cone damage rates, leading to a higher net production of sound seeds compared to mixed forests. This study indicates that ongoing changes in forest composition after land abandonment may impact tree reproduction.

     

X‐ray computed tomography reveals that intraspecific competition promotes soldier differentiation in a one‐piece nesting termite

Investment in soldier production in eusocial lineages involves a trade‐off between maintenance costs and defense benefits. Termites are eusocial insects that live in colonies organized into three castes: primary reproductives, soldiers, and workers or pseudergates. Neotermes chilensis (Blanchard) (Isoptera: Kalotermitidae) is a one‐piece nesting termite that nests and forages in a single piece of wood. Two scenarios may be of importance in a defense context of one‐piece nesting termites: during swarms, when colonies may be invaded by winged termites (alates) in search of a place to found a new colony, and when colonies of conspecifics are present within the same substrate. It was hypothesized that the ratio of soldiers to non‐soldiers would be higher at the onset of the swarming period and in substrates bearing more than one termite colony. A method based on X‐ray computed tomography (CT) was developed to study gallery connectivity in colonies of N. chilensis and caste composition within colonies. Computed tomography allowed the digital reconstruction of the galleries within the substrate, even when they belonged to different colonies, and was effective in distinguishing termites from substrate, and soldiers from reproductives and pseudergates. Using CT, the ratio of soldiers to non‐soldiers was shown to be highest in colonies within multicolonial scapes (i.e., neighboring colonies were present in the same substrate) during the swarming season, thus supporting our initial hypotheses. These results constitute a unique example of induced defenses arising from intraspecific interactions in termites.     

Impact of prairie fragment size on proportion of females and reproductive success of Lobelia spicata Lam., a gynodioecious species

• Due to ongoing human impacts, plant species increasingly occur in landscapes that are highly fragmented, with remaining natural habitats occupying small areas, resulting in populations that are smaller and more isolated than in previous time periods. This changed metapopulation structure is expected to have negative impacts on seed production. For example, the proportion of female plants within gynodioecious populations may be more volatile due to genetic drift in small populations associated with small habitat fragments, with concomitant impacts on seed production. My aims were to determine: (i) if variation in proportion of females is larger in smaller fragments; and (ii) if such changes in female frequency in small fragments result in reduced seed production.
• Thirty‐two populations of Lobelia spicata Lam., a gynodioecious species, were surveyed in 2000, 2001 and 2009 in the tallgrass prairie region of Midwestern North America (Illinois and Indiana, USA). Data were collected for: proportion of female plants, total number of flowering plants (measure of population size), seed set per plant and prairie fragment size (another measure of population size).
• The proportion of females is more variable in smaller prairie fragments. Seed number per fruit decreases as the proportion of females increases in a population, but only significantly for female plants. The number of flowering plants is positively associated with fruit production for both genders. Populations within larger prairie fragments have higher seed production.
• The reproductive consequences of habitat fragmentation depend on the plant breeding system. While both sexes were negatively impacted, females were more adversely affected.

     

Phorid fly parasitism rates vary between individual colonies of the leaf-cutting ant Atta laevigata

1. Phorid flies are important natural enemies of leaf-cutting ants. Female phorids lay eggs in the worker ants' bodies and the parasitoid larvae feed on the host's internal tissues.
2. Here we investigated the parasitism rates by four species of phorids when attacking individual Atta laevigata colonies at three different field sites in order to further understand the host-parasitoid relationship.
3. There were distinct variations in parasitism rates when comparing phorid species attacking individual colonies. In 13 of the colonies sampled, parasitism by Apocephalus attophilus or Eibesfeldtphora erthali was predominant. However, in four of the colonies, Myrmosicarius grandicornis and Apocephalus vicosae were the predominant species.
4. Variations in parasitism rates by the four phorid species were investigated in relation to the size of the potential hosts that had been collected from the foraging trails of each colony.
5. When correlating the size of potential hosts to phorid species, ants with head capsule widths of >2 mm were predominantly parasitized by A. vicosae and E. erthali. Smaller ants, with head capsule widths of 2 mm or less, were predominantly parasitized by A. attophilus and M. grandicornis.
6. The highest parasitism rates by E. erthali were observed in individual colonies that had the highest proportions of large workers on the trails.
7. Although almost half the size of E. erthali, A. attophilus parasitized both large (head capsule width > 2 mm) and small workers (<2 mm). This was related to the fact that between 1 and 16 larvae can develop within an individual host, depending on host size.
8. M. grandicornis parasitized smaller workers (mean head capsule width = 2 mm) as these phorids are themselves small and only one larva develops within each host.
9. This study demonstrated that the parasitism profiles of individual A. laevigata colonies were related to the size of foragers on the trails.

     

Losses of workers and reproductives in colonies of the paper waspPolistes riparius (Hymenoptera: Vespidae) due to the parasitic waspLatibulus sp.

1. Losses in workers and reproductives due to the ichneumonid parasitoid Latibulus sp. were determined in a haplometrotic, temperate paper wasp, Polistes riparius, during a 5-year study.
2. Fifty-four to 79% of pre-emergence nests were parasitized on worker brood, resulting in a worker loss of 25–31% on average. Worker brood reared in outer cells were much more often parasitized than those in central ones, possibly because outer cells are more accessible to the parasitoid. Infestation of worker brood was not random but aggregated among nests.
3. Seventy-eight to 100% of nests were parasitized on reproductive brood, and lost 10–34% of reproductives.
4. The number of emerged workers positively correlated with that of reproductives produced and that of cells made during the season. This suggests that worker loss reduces reproductive output of colonies. Hence, the parasitoid can reduce colonial reproductive output not only by killing reproductive brood but by reducing worker force.

     

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

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Alarm-recruitment behaviour in Pheidole militicida (Hymenoptera: Formicidae)

ABSTRACT.

• 1 Pheidole militicida Wheeler, a seed-harvesting species of the southwestern United States, possesses a major worker caste (soldiers) with unusually large heads. Previous work suggested that these large major workers are specialized defenders against large seed-harvesting species in the genus Pogonomyrmex.
• 2 Experimental introductions of Pogonomyrmex maricopa Wheeler and the omnivorous fire ant, Solenopsis xyloni McCook, demonstrate that P. militicida colonies alarm-recruit major workers against S.xyloni workers but not against the larger Pogonomyrmex workers.
• 3 P.militicida is the second species in which enemy-specific defence against Solenopsis fire ants has been detected. I suggest that the special role of Solenopsis as both competitor and predator in ant communities frequently favours major worker specialization against this genus.
• 4 Major workers of P.militicida and Pheidole dentata Mayr, the other species with enemy-specific defence against Solenopsis, are morphologically very different from one another. It is suggested that interspecific variation in major worker morphology is not necessarily associated with variation in behavioural specialization.