Behavioral evolution in the major worker subcaste of twig-nesting <Emphasis Type="Italic">Pheidole</Emphasis> (Hymenoptera: Formicidae): does morphological specialization influence task plasticity?

Polymorphism frequently correlates with specialized labor in social insects, but extreme morphologies may compromise behavioral flexibility and thus limit caste evolution. The ant genus Pheidole has dimorphic worker subcastes in which major workers appear limited due to their morphology to performing defensive or trophic functions, thus providing an ideal model to investigate specialization and plasticity. We examined worker morphology, brood-care flexibility, and subcaste ratio in 17 species of tropical twig-nesting Pheidole by quantifying nursing by major workers in natural colonies and in subcolonies lacking minors, in which we also measured brood survival and growth. Across species, majors performed significantly less brood care than minors in intact colonies, but increased rates of brood care 20-fold in subcolonies lacking minors. Brood nursed by majors had lower survival than brood tended by minors, although rates of brood growth did not vary between subcastes. Significant interspecific variation in rates of brood care by major workers did not lead to significant differences in brood growth or survival. Additionally, we did not find a significant association between the degree of major worker morphometric specialization and rates of nursing, growth, or survival of brood among species. Therefore, major workers showed reduced efficacy of brood care, but the degree of morphological specialization among species did not directly compromise task plasticity. The compact nests and all-or-nothing consequences of predation or disturbance on colony fitness may have influenced the evolution of major worker brood-care competency in twig-nesting Pheidole. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Dedicated to Professor Edward O. Wilson on the occasion of his 80th birthday.     

Provisional rejection of three alternative hypotheses on the maintenance of a size dichotomy in males of Dawson’s burrowing bee,Amegilla dawsoni (Apidae,Apinae, Anthophorini)

I examine three alternative hypotheses on the male size dimorphism of Dawson’s burrowing bees (Amegilla dawsoni) in which there are large (major) and small (minor) males. One possibility is that minor males are simply the incidental byproduct of environmental conditions that prevent females from provisioning brood cells optimally. This hypothesis is not supported by the finding that males of intermediate size are consistently rare in populations sampled across years and in different regions, nor can it easily account for the absence of a size dichotomy in females. A second possibility is that minors represent a “best of a bad job” response of those females that are small or otherwise disadvantaged. However, presumptive male siblings sometimes include both majors and minors, a result not predicted from this hypothesis. A third explanation is that female brood provisioning strategy results in the production of minors and majors with equal fitness benefit to fitness cost ratios. However, although it is true that minor males weigh on average about half what a major weighs, and so represent approximately half the provisioning expense of a major, minor males on average appear to secure far fewer than half the number of matings of majors. If the estimate of mating success of minors is accurate, the net gain to females from producing a minor son is unlikely to equal that derived from a major son. Therefore the third hypothesis must also be tentatively rejected, although with caution given the uncertainties in estimating the relative costs and benefits of producing major and minor sons. Received: 12 January 1996/Accepted after revision: 27 April 1996     

Nestmate recognition in three species of stenogastrine wasps (Hymenoptera,Vespidae)

The capacity to recognise a conspecific intruder was investigated in Parischnogaster jacobsoni, Liostenogaster flavolineata and L. vechti, three species of primitively social wasps of the subfamily Stenogastrinae. Results of behavioural experiments carried out in the field showed that females of all three species react pacifically if presented with female nestmates, but aggressively reject an intruder from a conspecific colony. As L. flavolineata and L. vechti both build large clusters of nests, often very close to each other, the recognition capacity among females from different nests, but in the same conspecific cluster, was also investigated. Females of both species were more aggressive towards females from a different colony in the same cluster than towards their female nestmates. Additional experiments on L. flavolineata showed that there was no difference in reaction towards females from colonies nearer or further from the tested colony but within the same cluster, nor towards females from a different cluster. The capacity to recognise an alien conspecific nest containing immature brood was investigated in P. jacobsoni. Adult females of this species, invited to land on an alien nest which had experimentally been exchanged for their own, accepted the new nest and partially destroyed the immature brood. The behaviour of the females when they land on an alien nest, however, suggests that they do recognise the nest as foreign. Acceptance of foreign nests coupled with low immature brood destruction is probably due to the high energetic costs of egg-deposition and larval rearing in stenogastrine wasps. These results suggest that nestmate recognition in these wasps is very efficient, even though they belong to the most primitive subfamily of social wasps. Received: 16 April 1996/Accepted after revision: 9 August 1996     

Age-related repertoire expansion and division of labor in Pheidole dentata (Hymenoptera: Formicidae): a new perspective on temporal polyethism and behavioral plasticity in ants

The controversy concerning the extent to which the organization of division of labor in social insects is a developmental process or is based on task allocation dynamics that emerge from colony need independent of worker age and endocrine or neural state has yet to be resolved. We present a novel analysis of temporal polyethism in the ant Pheidole dentata, demonstrating that task attendance by minor workers does not shift among spatially associated sets of behaviors that minimally overlap but rather expands with age. Our results show that the number of tasks performed by older minors increases through the addition and retention of behaviors, with up to a sixfold increase in repertoire size from day 1 to day 20 of adult life. We also show that older minors respond to colony needs by performing significantly more brood care as its demand increases, indicating that they can quickly upregulate nursing according to labor requirements. This level of plasticity was absent in younger siblings. The breadth of responsiveness to task-related olfactory stimuli increased with age. In a binary choice test in which young and old minor workers could orient toward odorants from brood or food, older workers responded to both brood and food, whereas young workers responded only to brood. These dissimilar responses to stimuli associated with nursing and foraging indicate age-related differences in sensory ability and provide a physiological basis for the age-related repertoire expansion model. We discuss repertoire expansion in P. dentata in light of behavioral development and caste flexibility in ants.     

Brood-care experience, nursing performance, and neural development in the ant Pheidole dentata

Social insect workers mature behaviorally and physiologically with increasing age, generally transitioning from or adding new tasks to their existing repertoire of within-nest nursing tasks. As adult minor workers of the ant Pheidole dentata age, they attend to brood more frequently and nurse more efficiently, perform a broader array of tasks, and undergo myological and neural development. Because these factors covary, the causal relationships among age, task experience, and neural and physiological maturation are not understood. We compared brood-care performance and efficiency by 10-day-old P. dentata minors that had acquired nursing experience to that of equal-age minors experimentally deprived of brood contact. We found the frequency and efficiency of nursing did not significantly differ between experimental and control worker groups, suggesting experience is not required for age-related improvement in nursing efficiency. Workers with and without prior nursing experience did not significantly differ macroscopically in brain anatomy or in brain serotonin content, although workers from the two treatments had slightly, but significantly, different levels of brain dopamine. These results suggest experience with brood is not required for P. dentata minor workers to develop nursing proficiency or undergo a substantial degree of the age-related neural development identifiable by our assessments, which could underscore the ontogeny of brood-care efficiency.     

Nest defense and early production of the major workers in the dimorphic ant Colobopsis nipponicus (Wheeler) (Hymenoptera : Formicidae)

Summary Production of the major subcaste and its contribution to nest survival in the dimorphic ant Colobopsis nipponicus was examined in the field. In this species, the first major workers were reared in the second brood, very early in the colony life cycle. A field experiment demonstrated that artificial colonies without major workers could not survive, whereas colonies with at least one major worker per nest entrance could. Because major workers of C. nipponicus defend the nest entrance by head plugging, the lack of nest defenders in the experimental colonies seemed to be a major cause of nest failure. The defensive value of major workers was much higher than that of minor workers. Many artificial colonies without major workers were displaced by competitors for nest sites, especially by those of other conspecific colonies. In addition, more than 90% of field colonies nested with other conspecific colonies on the same tree. The early production of major workers in C. nipponicus seemed to be very important for the survival of incipient colonies.     

The behaviour of ant transporters at the old and new nests during successive colony emigrations

Colonies of the ant Temnothorax albipennis improve their collective performance over successive emigrations (Langridge et al. Behav Ecol Sociobiol 56:523–529, 2004, Behav Ecol Sociobiol 62:447–456, 2008). Here, by analysing the performance of individual transporters (workers that carry the brood, queen and a proportion of adults), we investigate whether they spend less time at the old and new nests during repeated emigrations. Transporters expedited choosing and picking up brood items at the old nest and depositing them in the new nest. Such improvements were not associated with adult transport. Generally, when carrying brood items, but not when carrying adults, transporters visited several locations in the new nest before depositing them. Transporters did not interact with other adults when depositing brood items. Consequently, reductions in depositing times are the sum of time savings made by individual transporters. By contrast, transporters spent most time interacting with other adults before picking up brood items at the old nest. As the frequency of these interactions did not decline, we suggest the behaviours of interacting adults were modified in a way that hastened their completion. Thus, reductions in picking-up times probably occur because of time saved during interactions.     

Colony growth and the ontogeny of worker polymorphism in the fire ant,Solenopsis invicta

Summary Colony size and worker polymorphism (headwidth) were determined for fire ant colonies ranging from incipient to 12 years of age. Colonies grew approximately logistically, reaching half size between 21/2 and 31/2 yr and reaching their maximum size of about 220000 workers after 4 to 6 yr. Colony size showed strong seasonal variation. There was some evidence that growth rate may vary with food density. Incipient colonies are monomorphic and consist of small workers only, but as colonies grow, production of larger workers causes the size-frequency distributions to become strongly skewed. These skewed distributions were shown to consist of two slightly overlapping normal distributions, a narrow one defined as the minor workers, and a much broader one defined as the major workers. Major workers differ from minor workers in having been subjected to a discrete, additional stimulation of body growth, resulting in a second normal subpopulation. The category of media is seen to be developmentally undefined. The mean headwidth of the workers in both of these subpopulations increased during the first 6 mo. of colony life, until colonies averaged about 4000 workers. Headwidth of minors declined somewhat in colonies older than about 5 yr, but that of majors remained stable. When the first majors appear, their weight averages about twice that of minors. This increases to about 4 times at 6 mo. and remains stable thereafter. The range of weights of majors is up to 20 times that of minors. Growth of the subpopulation of major workers is also logistic, but more rapid than the colony as a whole, causing the proportion of major workers to increase with colony size. In full sized colonies, about 35% of the workers are majors. Total biomass investment in majors increases as long as colonies grow, beginning at about 10% at 2 months and reaching about 70% in mature colonies. This suggests that major workers play an important role in colony success. The total dry biomass of workers peaked at about 106 g, that of majors at about 72 g. These values then fluctuate seasonally in parallel to number of workers. When colony growth ceases, the proportion of majors remains approximately stable. Colony size explained 98% of the variation in the number of major workers.This is paper No. 18 of the Fire Ant Research Team     

Weak specialization of workers inside a bumble bee (<Emphasis Type="Italic">Bombus impatiens</Emphasis>) nest

Division of labor is common across social groups. In social insects, many studies focus on the differentiation of in-nest and foraging workers and/or the division of foraging tasks. Few studies have specifically examined how workers divide in-nest tasks. In the bumble bee, Bombus impatiens, we have shown previously that smaller workers are more likely to feed larvae and incubate brood, whereas larger workers are more likely to fan or guard the nest. Here, we show that in spite of this, B. impatiens workers generally perform multiple tasks throughout their life. The size of this task repertoire size does not depend on body size, nor does it change with age. Further, individuals were more likely to perform the task they had been performing on the previous day than any other task, a pattern most pronounced among individuals who guarded the nest. On the other hand, there was no predictable sequence of task switching. Because workers tend to remain in the same region of the nest over time, in-nest workers may concentrate on a particular task, or subset of tasks, inside that region. This division of space, then, may be an important mechanism that leads to this weak specialization among in-nest bumble bee workers.     

Brood sorting by ants: distributing the workload over the work-surface

Summary Leptothorax unifasciatus ant colonies occupy flat crevices in rocks in which their brood is kept in a single cluster. In artificial nests made from two glass plates sandwiched together, designed to mimic the general proportions of their nest sites in the field, such colonies arrange their brood in a distinct pattern. These patterns may influence the priority with which different brood are tended, and may therefore influence both the division of labour and colony demography. Different brood stages are arranged in concentric rings in a single cluster centred around the eggs and micro-larvae. Successively larger larvae are arranged in progressive bands away from the centre of the brood cluster. However, the largest and oldest brood items, the prepupae and pupae, are placed in an intermediate position between the largest and most peripheral larvae and the larvae of medium size. Dirichlet tessellations are used to analyze these patterns and show that the tile areas, the area closer to each item than its neighbours, allocated to each type of item increase with distance from the centre of the brood cluster. There is a significant positive correlation between such tile areas and the estimated metabolic rates of each type of brood item. The ants may be creating a domain of care around each brood item proportional to that item's needs. If nurse workers tend to move to the brood item whose tile they happen to be within when they have care to donate, they may apportion such care according to the needs of each type of brood. When colonies emigrate to new nests they rapidly recreate these characteristic brood patterns.     

Effects of foundress number on brood raids and queen survival in the fire ant Solenopsis invicta

In several ant species, colonies are founded by small groups of queens (pleometrosis), which coexist until the first workers eclose, after which all but one queen is killed. It has been hypothesized that, by producing a larger cohort of workers, cooperating queens may increase colony success during brood raids, a form of competition in which brood and workers from losing nests are absorbed into winning colonies. To test whether this benefit is sufficient to favor pleometrosis, newly mated queens of the fire ant Solenopsis invicta were assembled in groups of one, two, three, or four, reared in the laboratory until the first workers eclosed, then planted in the field in replicated assemblages. The proportion of colonies engaging in brood raids increased with average foundress number per nest and with colony density but was unaffected by variance in foundress number among interacting colonies. Within mixed assemblages of single-queen and multiple-queen colonies, queen number had no effect on the likelihood of engaging in raids or the probability of nest survival through the brood raiding period. However, following nearly 30% of raids, queens moved to new nests and displaced the resident queens. When queen relocation and subsequent mortality were accounted for, it was found that the survival of queens from four-queen groups was substantially higher than that of solitary queens. By contrast, the survival of queens from two-queen colonies was no greater than that of solitary queens. These results show that the competitive advantages of multiple-queen colonies are sufficient to counterbalance the increased mortality of queens within groups only when the number of foundresses is greater than two and when colonies are founded at high density. When colonies lose brood raids, the workers appear to abandon their mothers to join surviving colonies. However, in laboratory experiments, queens attempting to enter foreign nests were significantly more likely to displace the resident queen if their own daughters were present within the invaded nest. Thus, workers may be able to bias the probability that their mother rejoins them and displaces competing queens.     

Experimental analysis of worker division of labor in bumblebee nest thermoregulation (<Emphasis Type="Italic">Bombus huntii</Emphasis>, Hymenoptera: Apidae)

Bumblebee colonies experience daily and seasonal fluctuations in ambient temperature, but proper brood development requires a stable nest temperature. This study examined how adaptive colony responses to changing ambient temperature are achieved through the in-nest workers’ behavioral plasticity. We studied three Bombus huntii colonies in the laboratory. In the first experiment, we manipulated ambient temperature and recorded brood cell incubation and wing fanning by individually marked, known-age bees. The colonies maintained their nests closer to appropriate brood development temperatures (28 to 32°C) when exposed to a range of ambient temperatures from 10.3 to 38.6°C. Incubation activity was greater in cooler treatment conditions, whereas in the highest temperature treatment, some bees fanned and others moved off the brood. As the ambient temperature dropped, workers increased the duration of their incubating bouts, but, except at the highest temperature, the number of workers that incubated did not differ significantly among treatments. A subset of the bees incubated significantly more than their nest mates, some of which never incubated. Worker body size, but not age, was a good predictor of incubation rates, and smaller bees incubated at higher rates. In the second experiment, we removed the most actively incubating workers. Immediately after removals, the total colony incubation effort was lower than pre-removal levels, but incubation effort rebounded toward pre-removal levels after 24 h. The increased thermoregulatory demand after removals was met primarily by bees increasing their rates of incubation rather than by bees switching from a different task to incubation. We conclude that some B. huntii workers specialize on nest thermoregulation, and that changes in work rates are more important than task switching in meeting thermal challenges.     

Brood-provisioning strategies in Dawson's burrowing bee, Amegilla dawsoni (Hymenoptera: Anthophorini)

Males of Dawson's burrowing bee are dimorphic in size. Although large (major) males defeat smaller ones in competition for emerging females and therefore are more likely to mate, majors are greatly outnumbered by half-sized (minor) males. Nesting females might produce many minor males, despite their low reproductive value, because female behaviour is governed by a mixed evolutionarily stable strategy (ESS), in which case the ratio of majors to minors should not be affected by changes in female condition. In contrast, a conditional-strategy hypothesis predicts that older, wing-worn or stressed females unable to forage efficiently should be especially likely to produce minor offspring, which require less brood provisions. To test these alternative hypotheses, we manipulated the condition of nesting female bees by the addition of weights and the removal of their wing margins. These manipulations, done early in the flight season, failed to increase the production of minor males, a result consistent with the mixed-ESS hypothesis. However, unmanipulated females were far more likely to produce minor males if they were small or if they were nesting late in the season, when foraging conditions had deteriorated, findings that are consistent with a conditional provisioning strategy. Thus it appears that the abundance of minor males is the result of a conditional provisioning strategy of nesting females, which may be superimposed on a fixed tendency to produce large offspring early in the season and small ones later.     

Alternative mating tactics in dimorphic males of the harvestman <Emphasis Type="Italic">Longiperna concolor</Emphasis> (Arachnida: Opiliones)

Intense male–male competition for females may drive the evolution of male morphological dimorphism, which is frequently associated with alternative mating tactics. Using modern techniques for the detection of discontinuous allometries, we describe male dimorphism in the Neotropical harvestman Longiperna concolor, the males of which use their elongated, sexually dimorphic legs IV in fights for the possession of territories where females lay eggs. We also tested three predictions related to the existence of alternative mating tactics: (1) if individuals with relatively longer legs IV (majors) are more likely to monopolize access to reproductive resources, they are expected to remain close to stable groups of females more than individuals with relatively shorter legs IV (minors) do; (2) if minors achieve fertilization by moving between territories, they are expected to be less faithful to specific sites; and (3) majors should be observed in aggressive interactions more often. We individually marked all the individuals from a population of Longiperna during the reproductive season and recorded the location of each sighting for males and females as well as the identity of males involved in fights. Majors were more likely to have harems, and large majors were even more likely to do so. Majors were more philopatric and all males involved in fights belonged to this morph. These results strongly suggest that the mating tactic of the majors is based on resource defense whereas that of the minors probably relies on sneaking into the territories of the majors and furtively copulating with females.     

The function of polydomy: the ant <Emphasis Type="Italic">Crematogaster torosa</Emphasis> preferentially forms new nests near food sources and fortifies outstations

Many ant species are polydomous, forming multiple spatially segregated nests that exchange workers and brood. However, why polydomy occurs is still uncertain. We investigated whether colonies of Crematogaster torosa form new polydomous nests to better exploit temporally stable food resources. Specifically, we tested the effect of food presence or absence and distance on the likelihood that colonies would form a new nest. Because this species also forms little-known structures that house only workers without brood (outstations), we also compared the function of this structure with true nests. Laboratory-reared colonies were connected to a new foraging arena containing potential nest sites with or without food for 4 months. When food was present, most colonies formed polydomous nests nearby and the remainder formed outstations. When food was absent, the behavior of colonies differed significantly, frequently forming outstations but never polydomous nests. Distance had no effect on the type of structure formed, but when food was present, a larger proportion of the workforce moved shorter distances. Workers often fortified the entrances to both structures and used them for storage of dried insect tissue (“jerky”). In an investigation of spatial fidelity, we found that workers on the between-nest trail were associated with the original nest, whereas workers collecting food were more likely to be associated with the new nest or outstation. C. torosa appears to have a flexible colony structure, forming both outstations and polydomous nests. Polydomous nests in this species were associated with foraging and were only formed near food resources.     

The migrating herdsman Dolichoderus (Diabolus) cuspidatus: an ant with a novel mode of life

Summary The Malayan ant Dolichoderus cuspidatus lives in obligatory symbiosis with the pseudococcid Malaicoccus formicarii and other species of the same genus. The assemblies, which may be encountered up to 25 m away from the nest, are constantly covered with a great number of worker ants who protect them and receive honeydew. In the event of heavy rain the workers from a dense protective cluster, clinging to each other on top of the mealybugs. Neither hunting behavior nor active search for protein sources was observed in D. cuspidatus, although dead insects were accepted as food. When not searching for new plants, the activity of the ants outside the colony is limited to visiting the mealybugs. During the night and parts of the day the ants stay in their nest. Ant colonies deprived of their mealybugs are not viable due to their dependence on the symbiosis and because of the competition of other ants. Antless M. formicarii are likewise not viable. The mealybugs are extremely polyphagous and feed on many different monocotylous and dicotylous angiosperms. They feed exclusively on the phloem sap of young plant parts which are rich in amino acids. Dolichoderus cuspidatus workers carry the mealybugs to such locations. During the picking up and carrying process both partners display typical behavioral patterns. The colonization of new feeding sites takes place in well organized mass processions. During the foundation or disintegration of large feeding complexes, provisional depots with waiting mealybugs and ants are set up. The pseudococcids are carried not only while shifting the feeding sites, but also whenever the colony leaves its former nesting site and especially when any kind of disturbance occurs. They are even carried about without any apparent external cause, which leads to the fact that, at all times of trail activity, on average more than 10% of all ants using the trails carry mealybugs. Mealybugs are also present within the nest, especially adult females which are viviparous and give birth to their offspring there. Censused colonies each consisted of over 10 000 workers, about 4000 larvae and pupae, more than 5000 mealybugs and one ergatoid queen. Male winged ants were observed in large numbers during the dry season (January–February) and during the rainy season (September–October). The colonies form typical clumplike bivouac nests consisting of clusters of workers clinging to each other, thereby covering the brood and the mealybugs. The nesting site is in no way altered by constructive measures and is mostly found close to the ground. The preferred nesting sites are clusters of leaves, and cavities in wood or soil, although a freely hanging bivouac between a few branches may be set up as well. As soon as the distance between the nest and the feeding site is too great the colony moves to the feeding site, whereby the brood and the mealybugs are carried along in a well organized manner. During such nest-moving the establishment of intermediate depots can be observed. A shift of nest sites can also be induced by disturbances or by a change in the microclimate in the vicinity of the nest. Colonies multiply by budding. The tropical rain forest continuously offers different sprouting plants, the utilization of which requires extreme mobility on the part of the consumer. The unique behavioral strategy of D. cuspidatus, to carry constantly their polyphagous mealybug partners to new feeding sites and to take the whole colony there has enabled this ant and its symbiont to occupy this rich food niche. Dolichoderus cuspidatus is the first true nomad found in ants.     

Sanitizing the fortress: protection of ant brood and nest material by worker antibiotics

Social groups are at particular risk for parasite infection, which is heightened in eusocial insects by the low genetic diversity of individuals within a colony. To combat this, adult ants have evolved a suite of defenses to protect each other, including the production of antimicrobial secretions. However, it is the brood in a colony that are most vulnerable to parasites because their individual defenses are limited, and the nest material in which ants live is also likely to be prone to colonization by potential parasites. Here, we investigate in two ant species whether adult workers use their antimicrobial secretions not only to protect each other but also to sanitize the vulnerable brood and nest material. We find that, in both leaf-cutting ants and weaver ants, the survival of the brood was reduced and the sporulation of parasitic fungi from them increased, when the workers nursing them lacked functional antimicrobial-producing glands. This was the case for both larvae that were experimentally treated with a fungal parasite (Metarhizium) and control larvae which developed infections of an opportunistic fungal parasite (Aspergillus). Similarly, fungi were more likely to grow on the nest material of both ant species if the glands of attending workers were blocked. The results show that the defense of brood and sanitization of nest material are important functions of the antimicrobial secretions of adult ants and that ubiquitous, opportunistic fungi may be a more important driver of the evolution of these defenses than rarer, specialist parasites.     

Different policing rates of eggs laid by queenright and queenless anarchistic honey-bee workers (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Worker-reproduction is rare in queenright honey-bee colonies. When workers do lay eggs, their eggs are normally eaten by other workers presumably because they lack the queen's egg-marking signal. Workers use the absence of this queen signal to enforce the queen's reproductive monopoly by policing any worker-laid eggs. In contrast, in anarchistic colonies, the majority of the males arise from worker-laid eggs. Anarchistic worker-laid eggs escape policing because workers perceive anarchistic eggs as queen-laid. However, in this study, we show that eggs laid by queenless anarchistic workers do not escape policing and have very similar removal rates to worker-laid eggs from queenless wild-type (i.e. non-anarchistic) colonies. This suggests that, under queenless conditions, eggs laid by anarchistic workers lose their chemical protection and are therefore no longer perceived as queen-laid. Hence, the egg-marking signal seems to be only applied to eggs when queen and brood are present. This suggests that in the absence of queen and brood, the biosynthetic pathway that produces the egg-marking signal is switched off.Communicated by L. Keller     

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.