Solitary behavior in a high-altitude population of the social sweat bee Halictus rubicundus (Hymenoptera: Halictidae)

In the subalpine region of the Rocky Mountains of Colorado, United States, Halictus rubicundus has a solitary life cycle, but it is social in other parts of its known range. The brood is protandrous, with a nearly equal investment in the sexes. Productivity averages 6.5 offspring per foundress female, similar to the second brood of social nests in New York, but less than the combined productivity of both New York broods. Leucophora sp. (Diptera: Anthomyiidae) is the principal cause of brood mortality in Colorado. Foundress females in about half the nests survive until brood emerge as adults. Retention of these foundresses decreases offspring mortality by 68%. Comparable abilities to express solitary behavior with a single brood may characterize other eusocial halictine lineages that have successfully invaded high altitudes in the Rocky Mountains. The apparent inability to do this may help explain the absence of other eusocial halictine bees and polistine wasps at high altitudes, despite their success at lower elevations in the same mountains. Presence or absence of this ability may help explain latitudinal distributions of these lineages in North America. Holarctic distributions of lineages with eusocial behavior can be explained by migration as solitary populations from Eurasia to North America across Pleistocene Bering land bridges, with re-expression of double-brooded, eusocial behavior when the species then extended their ranges southward in North America. Received: 4 November 1994/Accepted after revision: 23 October 1995     

Caste determination and differential diapause within the first brood of Halictus rubicundus in New York (Hymenoptera: Halictidae)

Summary The partially bivoltine, primitively eusocial sweat bee Halictus rubicundus produces two female castes, gynes and non-gynes, in its first brood in New York. Castes in this brood differentiate within the first few days of adult life, with gynes leaving the population to overwinter as early as mid-June (non-gynes further differentiate into replacement queen and worker subcastes, with older females typically dominant). Analysis of possible mechanisms of caste determination reveals that although gynes average significantly larger than non-gynes within the first brood, this appears largely due to a late mean emergence data coupled with an increase in the size of emerging females over the course of brood emergence, rather than a causal relationship. A strong correspondence between male abundance (relative to newly-emerged females) and the pattern of gyne production, along with data from dissections, suggests that females that mate when young become diapausing gynes, while those that do not mate promptly become non-gynes and do not diapause even though many mate later. Although alternatives to this simple mechanism cannot be ruled out entirely, it nevertheless offers profound implications for theoretical and empirical understanding of the evolutionary origins of the worker caste.     

The role of learning in the mating biology of a sweat bee Lasioglossum zephyrum (Hymenoptera: Halictidae)

Summary Results from experiments on the role of learning in the mating biology of a sweat bee, Lasioglossum zephyrum (Hymenoptera: Halictidae), are described in this paper. Male learning of individual female odors is important in natural populations (Table 1), as has been shown in the laboratory. Four other hypotheses are unlikely to account for the observed behavior: (1) Female odors dissipate rapidly; (2) Males learn and then avoid the study area; (3) Males or females produce repellents which are effective against other males; or (4) males recognize their own odor on previously-contacted females, which they subsequently avoid. Regarding questions of male preference and optimal outbreeding, Tables 2 and 3 show there are no consistent preferences for more novel or less novel female odors.     

Trophic aspects of caste determination in Halictus ligatus,a primitively eusocial sweat bee

Caste determination in primitively eusocial sweat bees is thought to be due to an interacting suite of factors, including size of the larval provision mass, time of year, and social context of the nest into which a young female emerges. Newly emerged gynes are significantly fatter than newly emerged workers, suggesting the existence of larval caste determination cues. Since photoperiod, temperature, and interactions with nestmates were unlikely to affect larval caste determination, we compared the sizes and contents of larval provision masses destined to produce either workers or gynes. Gyne-destined larvae consumed pollen masses that were larger and contained slightly more sugar than those of worker-destined larvae. We suggest that sugar content is one cue which prompts the development of fat reserves in gyne-destined females but not in worker-destined females. The amount of fat possessed by a newly emerged female influences her chances of successfully entering diapause shortly after emergence. Therefore, small, lean females may be more susceptible to behavioural control by queens and more likely to become workers, while large, fat females would be more likely to become gynes. Correspondence to: M.H. Richards     

Solitary and eusocial nests in a population of Augochlorella striata (Provancher) (Hymenoptera; Halictidae) at the northern edge of its range

Summary Augochlorella striata was studied at the northern limit of its range. The study population contained a mixture of solitary and social nest foundresses. Eusocial foundresses produced 1 or 2 workers before switching to a male biased brood. Solitary foundresses produced males first. Cells vacated by eclosed offspring were reused late in summer. A female biased brood resulted from cell reuse in both solitary and eusocial nests. Workers were slightly smaller than their mothers and were sterile although most of them mated. In comparison to published data from a Kansas population of this species, the Nova Scotia population had i) a lower proportion of multiple foundress nests, ii) a smaller worker brood and iii) a briefer period of foraging activity but iv) comparable overall nest productivity.     

Assured fitness returns favor sociality in a mass-provisioning sweat bee,Megalopta genalis (Hymenoptera: Halictidae)

Assured fitness returns models for the evolution of sociality emphasize the selective value of ensuring that offspring receive adequate parental care to reach maturity. If a member of a social group dies, it can accrue returns on investment in offspring through the efforts of surviving social partners. We provide evidence that in the mass-provisioning, facultatively social sweat bee Megalopta genalis, adult presence in the nest throughout brood development provides protection from ant predation. Nests with adults present were well protected, and brood in nests with adults removed suffered higher predation. Females in observation nests showed effective defensive behavior against experimentally introduced ants, and bees in natural nests repulsed naturally occurring ant raids. Megalopta nest architecture and behavior are such that the brood of several cooperating females can be defended with little additional cost relative to solitary nesting. The benefits of cooperative defense may favor group living in mass provisioning bees. Our observations and experiments suggest that parental care throughout brood development can be adaptive in mass provisioning species, supporting the predictions of assured fitness returns models.     

Social evolution and its correlates in bees of the subgenus Evylaeus (Hymenoptera; Halictidae)

Summary Eleven behavioural characteristics of eight species of the subgenus Evylaeus were analysed using principal components analysis. The first component axis represents social level and explains over forty percent of the total variation in the data. The following characteristics are highly correlated with social level — (i) a reduction in the proportion of males in the first brood, (ii) a reduction in the proportion of workers that mate, (iii) a reduction in the proportion of workers that have developed ovaries, (iv) an increase in the mean number of workers, (v) increased contact between adults and developing brood and (vi) an increase in the size difference between queens and workers. Because these factors appear, in general, to be under the control of the queen it is argued that parental manipulation has been an important component of social evolution in these bees. The number of worker broods and the mechanism of male production are also related to social level but are less important. Nest architecture, nest defense and polygyny seem to be unrelated to social level. Variation in nest architecture may be in response to edaphic features of the substrate. The lack of any relationship between polygyny and social level implies that the semisocial route to eusociality was not the one taken. It is likely that polygyny can only occur where the substrate is suitable for the winter hibernation of sisters in their natal nest. Multivariate methods are useful in determining the relative social level of closely related halictine species.     

Attraction and learning in mate-finding by solitary bees,Lasioglossum (Dialictus) figueresi Wcislo and Nomia triangulifera Vachal (Hymenoptera : Halictidae)

Summary Males of the solitary sweat bee, Nomia triangulifera, patrol over large areas where thousands of females emerge, searching for receptive females. The daily operational sex ratio is strongly male-biased. Males contact dead, frozen (untreated) females more frequently than they contact females which were washed in hexane, showing that olfactory cues are utilized in mate-finding. A major source of female sex pheromone is in the head. Male pouncing on females is temporally non-random, indicative of group stimulation. Bioassays show that newly emerged females are more attractive to males than are older pollen-collecting females. Female odors are individually distinctive, based on male responses, and there is much variation among females in their attractive properties. Male responses to female odors suggest that learning is important for mating in natural populations. In contrast, the following hypotheses are unlikely to account for the observed behavior: (1) dissipation of female odors; (2) site learning and avoidance behavior by males; (3) decay of male motivation; or (4) male-produced repellents effective against other males. Laboratory and field studies show that female Lasioglossum figueresi produce individually-distinctive odors, which are attractive to males. There is considerable inter-individual variation among females in their attractiveness to males among sexually immature females. Male responses to female odors decay over the course of the presentation, suggesting the importance of learning in natural populations, although several alternatives could not be tested.     

Genetic population structure and sociogenetic organisation in Formica truncorum (Hymenoptera; Formicidae)

Summary The genetic population structure and the sociogenetic organization of the red wood ant Formica truncorum were compared in two populations with monogynous colonies and two populations with polygynous colonies. The genetic population structure was analysed by measuring allele frequency differences among local subsets of the main study populations. The analysis of sociogenetic organisation included estimates of nestmate queen and nestmate worker relatedness, effective number of queens, effective number of matings per queen, relatedness among male mates of nestmate queens and relatedness between queens and their male mates. The monogynous populations showed no differentiation between subpopulations, whereas there were significant allele frequency differences among the subpopulations in the polygynous population. Workers, queens and males showed the same genetical population structure. The relatedness among nestmate workers and among nestmate queens was identical in the polygynous societies. In three of the four populations there was a significant heterozygote excess among queens. The queens were related to their male mates in the polygynous population analysed, but not in the monogynous ones. The data suggest limited dispersal and partial intranidal mating in the populations with polygynous colonies and outbreeding in the populations having monogynous colonies. Polyandry was common in both population types; about 50% of the females had mated at least twice. The males contributed unequally to the progeny, one male fathering on average 75% of the offspring with double mating and 45–80% with three or more matings. Correspondence to: L. Sundström     

The chemical basis for trail recognition in Lasius nipponensis (Hymenoptera: Formicidae)

Summary. In this paper we report on the chemical basis for trail recognition in Lasius nipponensis. On and near trails in the field, workers became aggressive against conspecific intruders and succeeded in protective contests, while intruders usually avoided confrontation. Such asymmetric interactions were also observed in the laboratory when two non-nestmate workers were sequentially placed in a clean glass dish, while mutual aggression was observed when they were placed simultaneously. Asymmetric aggression was also observed when the workers were placed in a dish previously conditioned by other workers, or when the dish was treated with the hexane rinse of the conditioned dish. The rinse contained a series of hydrocarbons, in which components and proportion were almost identical to those of the cuticular hydrocarbons of the workers, except for a lack of n-alkanes. Amounts of the hydrocarbons that remained on the dish were very small when all the tarsi of the workers were incapacitated. We therefore consider that the hydrocarbons are not adhesion of the cuticular hydrocarbons but secretion from the tarsi of workers, and serve as a key signal for trail recognition in this ant.     

Predatory behavior of Polybia sericea (Olivier), a tropical social wasp (Hymenoptera: Vespidae)

Summary Polybia sericea (Olivier) (Hymenoptera: Vespidae) foragers were trained to visit experimental foraging plots and tests were conducted to determine the role of visual, olfactory, and chemotactile cues in prey location. Foragers approached prey from downwind and hovered downwind of visual and olfactory stimuli. Olfactory cues were more likely to elicit landing than were visual cues. Biting of potential prey was most consistently elicited by a combination of visual, tactile, and chemotactile cues. Foragers encountering large prey carried a piece back to the nest and returned for prey remains. These returning foragers used visual cues to direct intensive aerial search; olfactory prey cues elicited landing.     

Behaviorally mediated spatial and temporal refuges from a cleptoparasite,Argochrysis armilla (Hymenoptera: Chrysididae), attacking a ground-nesting wasp,Ammophila dysmica (Hymenoptera: Sphecidae)

Summary It has been suggested that parasite pressure favors the evolution of sociality within the Hymenoptera. I analyzed the impact of a chrysidid nest parasite, Argochrysis armilla, on its host, the solitary ground-nesting wasp, Ammophila dysmica, to assess the role of parasitism in favoring two steps towards sociality: aggregated nesting and delayed nest provisioning. The foraging strategy of Argochrysis armilla involves discovering host nests during excavation, learning the locations of discovered nest, and returning to oviposit in nests during nest provisioning; I therefore assessed the influence of host behavior on (1) parasite discovery of nests and (2) parasite oviposition in nests. Significantly fewer parasites discovered host nests that were excavated during the early morning and late afternoon, due to partial asynchrony of host and parasite activity. Nests excavated in areas of low nest density were also less likely to be discovered; use of low density nest sites increased during periods of high parasite activity. Due to a rapidly decelerating rate of parasite recruitment to nests under excavation, the duration of nest digging had only a limited influence on nest discovery by parasites. The probability of parasite oviposition in a host nest was determined by the number of parasites discovering the nest during excavation and by the time between nest excavation and provisioning; delaying nest provisioning reduced the risk of parasite oviposition. Delayed provisionings primarily appeared to be a result of the stochastic process of hunting and prey encounter. The number of provisions placed in a nest (one vs two) had no effect on the probability of nest parasitism. Spatial patterns of parasitism were directly density dependent in 1984 and density independent in 1986. In this system parasite pressure acts against the formation of nesting aggregations and in favor of delayed nest provisioning. The dependence of these results on species-specific aspects of the parasite's foraging strategy and the host's defensive strategy suggests, however, that different parasite species may generate qualitatively different selection pressures, potentially contributing to the diversity of nesting behavior in the Hymenoptera.     

Sex ratios and life-history patterns of a solitary wasp,Trypoxylon (Trypargilum) politum (Hymenoptera : Sphecidae)

Summary Werren and Charnov (1978) and Seger (1983) proposed a model to explain a fairly common pattern of alternating sex ratio biases between generations in partially bivoltine insects. When first-generation males overlap and mate with females of the second generation, then females should bias sex ratios in favor of sons for the first generation and daughters for the second generation. In an intensive, 7-year study at four sites in northern Florida, pipe-organ mud-daubing wasps (Trypoxylon (Trypargilum) politum; Hymenoptera: Sphecidae) were found to have strongly male-biased sex ratios in the first or overwintering generation and 1:1 or female-biased sex ratios in the second or summer generation. These differences were not due to differences in mortality of the two sexes but rather resulted from changing female sex-allocation decisions. In some respects the mud dauber results fit Seger's model well: alternating sex ratios in partially bivoltine populations, first-generation males overlapping second-generation females and perhaps most convincingly, northern, univoltine populations do not have a male bias. Despite this qualitative fit, however, our data do not meet the quantitative predictions of the model. This could result from the fact that some assumptions of the model are not met by the life history of T. politum. Alternative explanations for alternating sex ratios include split sex ratios, seasonal differences in cost ratios, facultative maternal investment rules and facultative overwintering decisions by offspring. Despite the position that sex ratios have achieved in the modern study of evolution, it is clear that accurate, quantitative predictions on sex-allocation patterns demand the same detailed understanding of the biology of the organism that is required for the study of other adaptations.     

The morphology and behavior of dimorphic males in Perdita portalis (Hymenoptera : Andrenidae)

Summary In Perdita portalis, a ground nesting, communal bee, males are clearly dimorphic. The two male morphs are easily distinguished based on head size and shape into (1) a flight-capable, small-headed (SH) morph that resembles the males of other closely related species and (2) a flightless, large-headed (LH) morph that possesses numerous derived traits, such as reduced compound eyes, enlarged facial foveae and fully atrophied indirect flight muscles. The SH morph occurs exclusively on flowers while the LH morph is found only in nests with females. While on flowers, SH males are aggressive, fighting with conspecific males and heterospecific male and female bees, and they mate frequently with foraging females. Using artificial observation nests placed in the field, I observed the behavior of females and LH males within their subterranean nests. LH males are aggressive fighters; males attacked each other with mandibles agape, and male-male fights always ended in the death of one male. LH males are highly attentive to the reproductive behavior of females; they spend increasing amounts of time near open cells during cell provisioning, and mating only takes place immediately prior to oviposition when females are forming the accumulated pollen and nectar into a ball. Based on larvae reared to adulthood in the laboratory, the two male morphs occur in equal proportions. The behavior of males in closely related species, especially P. texana, and the origin and maintenance of male dimorphism are discussed.     

Trail discrimination signal of Lasius japonicus (Hymenoptera: Formicidae)

Summary. Trail-following behavior of Lasius japonicus was colony-specific in the field, while trail pheromone activity was not. We found that the footprint substance caused colony-specific trail-following behavior only when working in conjunction with the trail pheromone. The footprint substance alone did not lead the workers to follow trails. The substance consisted mainly of hydrocarbons with composition almost identical to that of cuticular hydrocarbons, except for the absence of n-alkanes. Nestmate workers shared footprint hydrocarbon profiles as well as cuticular hydrocarbons, but the profiles differed among colonies. We therefore consider that the footprint hydrocarbon profiles serve as the trail discrimination signal in L. japonicus.