Colony Social Organisation and Alternative Social Strategies in the Eastern Carpenter Bee, <Emphasis Type="Italic">Xylocopa virginica</Emphasis>

The eastern carpenter bee, Xylocopa virginica, can nest either solitarily or in groups. In southern Ontario, Canada, near the northern edge of the range, most nests are social, containing groups of two to five adult females. Although social nests were much more frequent than solitary ones, they produced no more brood, so per capita brood productivity was actually lower for social females. Social females exhibited several reproductive strategies inferred from wing wear patterns, which reflect flight activity, and mandibular wear patterns, which reflect nest construction activity. Primary females accumulated a large degree of wing and mandibular wear and were presumed to be the primary reproductives in social nests. Secondary females accumulated less wing and mandibular wear and were probably subordinates awaiting opportunities to supersede primaries as dominant foragers and egg-layers. Tertiary females remained inactive, apparently deferring reproduction to the subsequent year. Social nesting, serial replacement of dominant females in social nests, and deferred reproduction are probably responses to severe competition for nests and nesting substrate.     

Breeding seasonality among long-tailed macaques,Macaca fascicularis,in Peninsular Malaysia

The females among more than 7000 long-tailed macaques, Macaca fascicularis,that had recently been trapped for biomedical research were examined for pregnancy. The animals were taken from the wild in the north of Peninsular Malaysia during the period 1978 to 1979 and in the south from 1978 to 1982. Both samples showed some pregnancies in all months of the year and a peak during the months of May to July. Additional peaks in other months in some years showed that, while the midyear pregnancy peak was significant, it was not as marked a phenomenon as is the breeding peak in many other species. This might be due to the complexity and unpredictability of both the climate and the macaques’ food supply in Peninsular Malaysia. The additional peaks also illustrated the pitfalls of generalizing about breeding seasonality on the basis of small samples.     

Ecological constraints on independent nesting in facultatively eusocial hover wasps

Recent field experiments suggest that cooperative breeding in vertebrates can be driven by a shortage of breeding territories. We did analogous experiments on facultatively eusocial hover wasps (Stenogastrinae: Liostenogaster flavolineata). We provided nesting opportunities by removing residents from 39 nests within a large aggregation (1995), and by glueing 20 nests obtained from a distant site into a second aggregation (1996). We prevented nest-less floaters from competing for these opportunities in 1995 but not in 1996. In both years, helpers in unmanipulated groups were given opportunities to nest independently without having to incur nest-building costs and with a reduced wait before potential helpers emerged. Helpers visited the nests we provided, but adopted only a small proportion (5% of 111 vacancies created in 1995). Others were adopted by floaters, but a significant proportion of nests were never adopted (nine out of 20 in 1995, seven out of 20 in 1996). Helpers that visited nests did not originate from particular kinds of social group. Nests containing older brood were more likely to be adopted, and adopting females rarely destroyed older brood. A general feature of social insect but not vertebrate life-histories, the long period of offspring dependency relative to the short life expectancy of adult carers, may be a key factor constraining independent nesting.     

Colony Development and Serial Polygyny in the Primitively Eusocial Wasp Belonogaster juncea juncea (Vespidae: Polistinae)

In Cameroon, the social ropalidiine wasp Belonogaster juncea juncea mostly selects human constructions for nesting, but nests installed in the shelter of large boulders were also noted. Nests were founded throughout the year by one female (25.5%) or by associations of two to eight females. Among the 49 colonies studied from foundation, 24 produced at least one adult, and only 5 produced males among the eight that succeeded in producing sexual individuals. After the first dominant female disappeared (died or abandoned the nest, from 77 to 196 days after foundation) the remaining females fought among themselves, leading to a drastic decrease in the total number of brood. They completely abandoned 22 of 24 surviving nests, over a mean colony cycle lasting about 7 months. Hence the colony and nesting cycles were congruent, illustrating that determinate nesting corresponds to the majority of the cases in this species. In the other two nests, some females remained on the nest and began a new colonial cycle illustrating semi-independence between colony and nesting cycles. Up to four successive cycles were completed in this way in the same nest, thus illustrating the phenomenon of serial polygyny (intermediate between determinate and indeterminate nesting cycles). Serial polygyny in independent founding polistine wasps, previously documented only once, has implications for studies attempting to identify factors involved in regulating the colony and nesting cycles in tropical social wasps.     

Sociality in the phylogenetically basal allodapine bee genus Macrogalea (Apidae: Xylocopinae): implications for social evolution in the tribe Allodapini

Recent phylogenetic studies based on DNA sequence data indicate that the tropical African bee genus Macrogalea is the sister group to the remaining extant allodapine fauna, whereas previously it was thought to be a distal group. This leads to some fundamental changes in our understanding of social evolution in the allodapines. Earlier studies suggested that Macrogalea showed only weak forms of social behaviour and these were not well characterized. However, large samples of Macrogalea zanzibarica presented here show that this species exhibits marked social behaviour. Nearly half of nests collected contained two or more adult females, with up to 10 females per nest. Brood are reared progressively and brood ages within colonies are staggered, giving rise to colonies with very mixed age structures and therefore frequent opportunities for alloparental care. Ovarian dissections indicate non‐simple forms of reproductive partitioning within colonies and most multi‐female colonies show evidence that more than one female has contributed to egg production, though reproductive episodes among colony members are frequently asynchronous. Some females show signs of much higher wing wear than their nestmates, but always show signs of previous reproduction. Reproductive division of labour appears to be temporally marked, ovarian differentiation among nestmates is linked to relative body size, but permanent worker‐like or queen‐like castes appear to be absent. This is similar to the communal, continuously brooded and multivoltine behaviour of some tropical halictine species and may be due to the aseasonal nature of brood development in tropical regions. Patterns of per capita brood production indicate large benefits to multi‐female nest occupancy, and sex allocation is strongly female biased. These findings strongly suggest that the capacity for complex social interactions and alloparental care is an ancestral trait for all of the extant allodapine lineages. Therefore comparisons among extant allodapines are unlikely to throw light on the initial origin of social behaviour, though they may uncover origins of true caste behaviour and reversals to solitary nesting. Sex ratios in Macrogalea and most other allodapine genera, spanning a broad phylogenetic and ecological range, suggest that one or more allodapine traits have provided persistent selection for female‐biased sex allocation.     

Extreme Polygyny: Multi-seasonal “Hypergynous” Nesting in the Introduced Paper Wasp <Emphasis Type="Italic">Polistes dominulus</Emphasis>

In temperate climates, female paper wasps typically initiate new colonies in the spring. Several nest-founding tactics have been documented in Polistes species, including solitary nest initiation, joining a cooperative association, usurping an existing nest, or adopting an abandoned nest. Occasionally, exceptionally large groups of females have also been found reusing nests from the previous season. Here we report this phenomenon in introduced populations of the Eurasian species Polistes dominulus. We describe in detail the demographic and genetic characteristics of one such spring colony from Los Angeles, California, USA, which was collected with 84 associated adults and all stages of developing brood in its 613 cells. Genetic and morphological data indicate the presence of multiple reproductively active females of varying relatedness, as well as many nonbreeding females, including probable early-produced offspring. Despite some evidence of chaotic social conditions, the colony appeared to have been highly productive. Additional observations of similar colonies are needed to determine how control is maintained within such a large breeding aggregation.     

Simultaneous spawning by female stream goby Rhinogobius sp. and the association with brood cannibalism by nesting males

A laboratory experiment was conducted by varying the undersurface area of nesting substratum and the number of females in an experimental tank to elucidate the determinants of the mating pattern in the stream goby, Rhinogobius sp. cross‐band type. Males with larger nests tended to attract two or more females to their nest in a tank. Moreover, males spawned simultaneously with multiple females and entire brood cannibalism by males was rarely observed under a female‐biased sex ratio. When males spawned with a single female with low fecundity, however, entire brood cannibalism occurred at a high frequency, suggesting that a male guarding a nest with fewer eggs consumes the brood. Therefore, spawning behaviour of females that leads to a large egg mass would decrease the risk of entire brood cannibalism. In this species, simultaneous spawning by multiple females in a nest serves as a female counter‐measure against entire brood cannibalism. These results suggest that a conflict of interest between the sexes through brood cannibalism is a major determinant of simultaneous spawning.     

Social Behaviour of the Carpenter Bee Xylocopa pubescens (Spinola)

The development of about 20 relatively small nests of Xylocopa pubescens was studied. After the first offspring had become adult, these nests reached a social stage in which there was only one egg-layer per nest. Freshly emerged (teneral) adults eat a lot of food, collected by a forager, before they fly out of the nest. This food appears to be of major importance to these bees in that it makes them fully agile. It seems therefore, that part of the food needed for the development of a young bee is not given at the larval stage, via the beebread, but at the teneral adult stage. As a result of this, a necessary overlap of generations is accomplished. Besides, less pollen has to be collected for the provision of a brood cell, so more cells can be constructed within a short period of time. There is a high degree of nest competition. Many nests were taken over by conspecific individuals (or by females of X. sulcatipes) that were searching for nesting sites within the study area. However, although more than 50% of the solitary nests were taken over sooner or later, strangers hardly ever intruded into social nests, with more than one adult. This illustrates how important it is for a reproducing female to tolerate the presence of nestmates which guard the nest in her absence. Although in most cases the foundress of a nest proceeded to produce new brood in the presence of her offspring, it happened that a daughter took over reproduction from her mother. In two of these cases, it could be observed that a mother started a new nest elsewhere after having been thrown out by her own daughter. At least in these cases, nest competition between mother and daughter started long before the mother had reached the end of her reproductive capacities. Since nesting possibilities are scarce, it seems a logical strategy to stay in the maternal nest and wait for a chance to become the egg-layer if the mother dies or if she loses her dominance. The occurrence of several social interactions among nestmates is discussed: — trophallaxis was observed often, not only under ‘forced’ conditions, but also as a form of ‘voluntary’ feeding; — nestmates were observed to groom each other.     

Group nesting and individual variation in behavior and physiology in the orchid bee <Emphasis Type="Italic">Euglossa nigropilosa</Emphasis> Moure (Hymenoptera,Apidae)

The Euglossini are a key group for studying the traits that promote or hinder highly social behavior in bees because it is the only tribe in the Apine clade without large colonies or females with distinct life histories, e.g. queens and workers. There have been few studies on behavior of orchid bee females in nests because these nests are not found easily. Taking advantage of the relatively high abundance of Eg. nigropilosa nests at Reserva Natural La Planada, Colombia, we examined social behavior of Eg. nigropilosa individuals in five nests (3 original and 2 reused) for nine months. We report this species to have the largest colonies known for Euglossa, with nests reaching up to 22 individuals, and all nests containing more than one female bee from the same generation. These nests presented many traits that correspond to communal insect colonies. No generational overlap and no cooperative brood care were detected. We examined natural enemies and resource limitation as important factors for group nesting. We examined parasitoid attacks to cells in a nest with females and one without females. We also searched for nesting locations and examined nest re-use as indicators of nest site limitation. Lastly, we examined behavioral and physiological differences among females in the same nest. Such differences could be the bases for evolution of alternative life histories among group living females. We examined extent of ovary development and oviposition rates in similarly aged females in the same nest. We found large variation in reproductive effort of young females. We also examined differences in resin foraging and cell usurpation behaviors. Behavioral specialization was observed, with some individuals bringing only resin to the nest. Inside the nests, bees had territories in which they constructed and defended cells. This territoriality may be a defense against usurpation of provisioned cells by nest mates. Received 10 December 2007; revised 2 May 2008; accepted 7 May 2008.     

FEMALE-BIASED SEX RATIOS IN A FACULTATIVELY SOCIAL BEE AND THEIR IMPLICATIONS FOR SOCIAL EVOLUTION

Montane populations of the Australian allodapine bee, Exoneura bicolor, are characterized by high levels of cooperative nesting and strongly female-biased sex ratios. A conspecific population from heathland shows much lower levels of cooperative nesting and lower levels of female bias. In both habitats, sex-ratio bias is greatest in the smallest brood sizes and becomes successively less biased in larger broods. Parity is approached in the largest heathland colonies, but not for any brood-size category in montane areas. Adult intracolony relatedness is moderately high for colonies in both reused and newly founded nests in the montane habitat, but probably low or zero for newly founded nests in heathland. Colony efficiency, measured as the number of brood per adult, increases with colony size in both habitats, suggesting that cooperation between females increases mean female fitness. It is argued that patterns of sex allocation are consistent with nonlinear fitness-return models, in which the mean reproductive value of daughters increases with the number of daughters produced in a brood. Such increases probably arise from a number of social interactions, including cooperative brood defense, increased task efficiency, and lower per capita costs in nest construction. The term “local fitness enhancement” is introduced here to describe these effects collectively. The female-biased ratios should lower selective thresholds for sib-directed altruism, at least in the earlier stages of colony development. It is argued that local fitness enhancement facilitates eusociality in allodapine bees and could also play a role in other haplodiploid taxa, provided cooperative nesting largely involves sisters, colony efficiency increases with colony size, and optimal colony sizes are only achieved after two or more generations after founding.     

Rank and colony defense against conspecifics in a facultatively eusocial hover wasp

An important benefit of social living is increased capacityfor defense. Highly eusocial species have often evolved a fightingcaste for this purpose, but many facultatively eusocial insectsand cooperatively breeding vertebrates lack morphological castesand the decision to defend or not can depend on costs and benefitsto each individual. Defense by subordinates in a social groupcan be regarded as a form of helping, and helping input oftenvaries among subordinates of different age or size. Severalhypotheses attempt to explain variation in helping effort, includingthe effects of relatedness and differences in the costs of helping.Evidence for these hypotheses is mixed and often lacks dataon the rank of subordinates, an important determinant of expectedfuture fitness. We examined individual variation in propensityto defend the nest against conspecifics in the tropical hairy-facedhover wasp Liostenogaster flavolineata. Prior to experimentation,we determined the positions of all wasps in the age-based queueto inherit the single egg-laying position in each L. flavolineatagroup. Two approaches were then used: observations of defenseagainst natural intrusions by conspecifics and experimentaltrials where wasps were presented attached to a wire. Higherranks were more likely to defend the nest than lower ranks,opposite to the pattern previously documented for another formof helping: foraging effort. Possible explanations for thisresult are that higher ranked females are better defenders andthat they suffer a larger decrease in expected future fitnesswhen an intruder usurps their position in the inheritance queue.     

Major benefits of guarding behavior in subsocial bees: implications for social evolution

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.     

Pupal developmental temperature and behavioral specialization of honeybee workers (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Honeybees (Apis mellifera) are able to regulate the brood nest temperatures within a narrow range between 32 and 36°C. Yet this small variation in brood temperature is sufficient to cause significant differences in the behavior of adult bees. To study the consequences of variation in pupal developmental temperature we raised honeybee brood under controlled temperature conditions (32, 34.5, 36°C) and individually marked more than 4,400 bees, after emergence. We analyzed dancing, undertaking behavior, the age of first foraging flight, and forager task specialization of these workers. Animals raised under higher temperatures showed an increased probability to dance, foraged earlier in life, and were more often engaged in undertaking. Since the temperature profile in the brood nest may be an emergent property of the whole colony, we discuss how pupal developmental temperature can affect the overall organization of division of labor among the individuals in a self-organized process.     

How can cleptoparasitic drosophilid flies emerge from the closed brood cells of the red Mason bee?

Aculeate Hymenoptera provision their progeny with large amounts of food. To protect their investment against brood parasites, females of many bee and wasp species construct brood cells that are hard to penetrate when finally sealed. However, the sealed brood cells also pose a problem for parasites that oviposit in the brood cell during provisioning. Brood parasites are smaller than their host and may lack strong mandibles to break through the solid brood cell walls. Furthermore, in nests built in existing cavities, newly‐eclosed brood parasites need information about the location of the nest entrance. In the present study, the mechanisms of emergence are investigated in Cacoxenus indagator Loew (Diptera, Drosophilidae), the major cleptoparasite in nests of the red Mason bee Osmia bicornis L. (Hymenoptera, Megachilidae). Larvae of C. indagator move to brood cells closer to the nest entrance and sometimes make small emergence holes in the final closure of the nest entrance. Nevertheless, approximately one‐third of newly‐eclosed flies orientate and break through at least one intact cell partition to emerge. Flies make most of their attempts to emerge at the correct side (i.e. the one pointing to the nest entrance, probably by using the shape of the cell partition as a cue). Newly‐eclosed flies use their head blister (ptilinum) to exert hydraulic pressure on particles of the cell partitions and produce small holes. Thus, C. indagator exhibits a set of behavioural and physiological adaptations enabling them to successfully emerge even from closed brood cells of their host.     

Sociality in <Emphasis Type="Italic">Euglossa</Emphasis> (<Emphasis Type="Italic">Euglossa</Emphasis>) <Emphasis Type="Italic">viridissima</Emphasis> Friese (Hymenoptera,Apidae, Euglossini)

Euglossa viridissima is an orchid bee that forms both solitary and multiple female nests, making it a suitable species for the study of factors leading to diverse degrees of sociality in Euglossines. We conducted observations in eight reused nests (where a first generation of bees had been produced) kept in artificial boxes from the Yucatan Peninsula, Mexico. Five nests were reused (reactivated) by a single female (SFN), two nests reused by a mother and one daughter (MFN1) and one nest reused by the mother and two daughters (MFN2). No single nest was reactivated by unrelated females. The number of foraging trips, their duration and the duration of cell provisioning was not different between SFN and MFN. The overall production of cells per female was not different either between both types of nest. However, in MFN although all females did lay eggs, there was a reproductive skew in favor of the mother (95 and 45% of the brood produced in MFN1 and MFN2 respectively). She showed reproductive control of her daughters through oophagy and displaying threatening behavior when the daughters tried to open a cell where she had laid an egg. Brood losses to parasites (Anthrax sp. (Bom-byliidae) and Hoplostelis bivittata (Megachilidae)) were only found in SFN which possibly reflects and advantage of MFN in this respect. Our results coupled with other studies in Euglossa, reveal that a wide range of social behaviors occur in this genus, from solitary and communal to primitive reproductive division of labor. Multiple factors involving different levels of pressure imposed by food availability and parasites may favor such a diverse range of nesting behaviors. Interestingly, female associations in E. viridissima seem a result of kin selection that is enforced by coercion from mother females on their daughters. More studies are needed to shed light upon the social organization of Euglossa and other Euglossines and on their phylogenetic relationships in order to trace the origins of eusociality in Apidae. Received 12 February 2008; revised 25 June 2008; accepted 17 July 2008.     

Common Cuckoos Cuculus canorus change their nest‐searching strategy according to the number of available host nests

In recent decades, numerous studies have examined factors affecting risk of host nest parasitism in well‐known avian host–parasite systems; however, little attention has been paid to the role of host nest availability. In accordance with other studies, we found that nest visibility, reed density and timing of breeding predicted brood parasitism of Great Reed Warblers Acrocephalus arundinaceus by the Common Cuckoo Cuculus canorus. More interestingly, hosts had a greater chance of escaping brood parasitism if nesting was synchronized. Cuckoo nest searching was governed primarily by nest visibility at high host‐nest density. However, even well‐concealed nests were likely to be parasitized during periods when just a few hosts were laying eggs, suggesting that Cuckoos adjust their nest‐searching strategy in relation to the availability of host nests. Our results demonstrate that host vulnerability to brood parasitism varies temporally and that Cuckoo females are able to optimize their nest‐searching strategy. Moreover, our study indicated that Cuckoos always manage to find at least some nests to parasitize. Thus, in this case, the co‐evolutionary arms race should take place mainly in the form of parasitic egg rejection rather than via frontline pre‐parasitism defence.     

Reconstruction ofRopalidia fasciata nests by progeny females after a typhoon and its significance in the social evolution of wasps

A strong typhoon, Typhoon 13, which swept through Okinawa on 2 and 3 September 1993, knocked down 57 to 61 of 97Ropalidia fasciata nests. Thirty-five colonies soon reconstructed their nests, and these nests produced 54±46 adults per nest during the remaining two and a half months before winter. The number of females involved in nest reconstruction was 21.3±15.7 per nest. The frequency of foraging activity was significantly higher in nests being reconstructed than in normal nests. As only one out of 205 foundresses marked in spring was found in early September, most cases of nest reconstruction were considered to be made exclusively by females that emerged during the spring and summer of 1993 (progeny females). All of the reconstructed nests but one produced adult females. Many females (31%) collected from the two reconstructed nests were inseminated. As all the reconstructed nests became vacant by early January, indicating completion of the colony cycle, and a large number of nests were established in the spring of 1994, females emerging from reconstructed nests in 1993 probably become foundresses in 1994.Ropalidia fasciata is considered to be, at least partially, bivoltine in Okinawa. The significance of these facts for the evolution of multi-queen social systems in the Polistinae is discussed.     

Sulcopolistes atrimandibularis,Social Parasite and Predator of an Alpine Polistes (Hymenoptera,Vespidae)1

Sulcopolistes atrimandibularis Zimmermann is the obligate social parasite of Polistes biglumis bimaculatus Geoffry in Furcroy, a mountain species of paper wasp. Unlike all the other hymenopteran social parasites, the Sulcopolistes female obtains part of the food for her immature brood by plundering other nests of the host species. Parasite females can control more than one host nest: one of them (the nursery nest) she uses solely for reproduction purposes (from which the Sulcopolistes reproductives emerge) and others (supply nests) are used for exploiting the Polistes brood content. It is possible that this behaviour is an adaptation to the extremely short colonial cycle of the host.     

The role of young guards inXylocopa pubescens

Summary Xylocopa pubescens is a facultatively social species in which two types of guards can be found: 1) old, formerly reproductive guards and 2) young, pre-reproductive guards that usually guard the nest in which they emerged. In this species it is always the dominant female that forages and lays the eggs.This paper focuses on the young females' reasons for guarding. Young guards are characteristically the first females of an emerging brood. They start guarding at an age of 6 days, and continue to do so for on average 10 days, when they start a dominance contest. In comparison to other emerging bees, guards did not receive any more of the incoming food. The presence of young females in the nest was sufficient to deter pollen robbers; the protection was not ameliorated by guarding. Guards did not protect the nest from usurpation by intruding females. The presence of a young guard positively influenced both the number and the duration of foraging flights.All guards attempted to take over dominance inside the nest by fighting with the dominant female. Guards had a probability of 50% of winning this contest, which was distinctly higher than their probability of finding a nest elsewhere. The dominant female was not in all cases the mother of the guard. Therefore, the average increase of indirect fitness by guarding was lower than the expected direct fitness returns from leaving the nest earlier. We therefore conclude that guarding females are environmentally disabled, hopeful reproductives. They may be guarding to determine the right time to risk a fight about dominance and to increase their direct fitness if their attempt to supersede is successful.