Process of comb division of reused nests in the Australian paper waspRopalidia plebeiana (Hymenoptera,Vespidae)

Summary InRopalidia plebeiana, combs made in the previous year are often reused by foundresses in the following spring and structurally divided into subnests by them. Close observations of the comb dividing process revealed that: 1) combs that had multiple brood areas (areas where cells had eggs) were frequently divided into subnests, while those with single brood areas were never divided; 2) groups of foundresses each occupied a particular brood area even before comb division started; and 3) frequency with which a foundress practiced comb cutting was independent of her social status or oviposition frequency. These observations suggest that the initial grouping of foundresses that colonize an old nest leads to subsequent comb division, rather than the major egg layers cutting combs to safeguard their oviposition territories from their rivals.     

Colony Foundation and Nestmate Recognition in the Social Wasp,Belonogaster petiolata

Emergence from hibernation and synchronized nest foundation by foundresses of B. petiolata in the southern Transvaal, appeared to be stimulated largely by a rise in temperature during early and mid August of each season. Initial foundresses on newly built nests were joined by other foundresses during the first few weeks of the nesting season to form foundress associations. Although females were strongly philopatric and able to discriminate between nestmates (from their natal nests) and non-nestmates, former nestmates did not reassociate to a significant degree on new nests. Thus, associated foundresses were, on average, not closely related. Rather than facilitating the reassociation of former nestmates, the advantage of philopatric behaviour appears to lie in ensuring that foundresses return to an area of relatively high nesting density, where their chances of being involved in multiple-foundress groups (irrespective of intra-colonial relatedness) are high. Non-philopatric females which nest away from localities of high nesting density have a low probability of being involved in associations, and may therefore have to attempt colony foundation alone, with little or no chance of reproducing successfully.     

Lipid stores, ovary development, and brain gene expression in Polistes metricus females

In order to gain insights into the mechanistic basis of caste and behavioral differences in Polistes paper wasps, we examined abdominal lipid stores and ovary development in Polistes metricus females in four groups: foundresses, queens, workers, and gynes. Queens had the largest ovaries, followed by foundresses, workers, and gynes. Gynes had 6x higher lipid stores than the other groups, and lipid stores were lower in foragers (foundresses, workers) than non-foragers (queens, gynes). Lipid levels and ovary development were negatively correlated across the four groups, but removing gynes from the analysis revealed a significant positive correlation for foundresses, workers, and queens, suggesting different energy allocation strategies for gynes vs. other groups. Expression levels of 9 genes (including three in the insulin pathway), examined in a previous study, correlated with either lipid stores or ovary development. These correlative results suggest important relationships between nutrition, reproduction, and division of labor in primitively social insects. We also show that it is possible to assign P. metricus females to one of the four female groups on the basis of wing wear (an indicator of foraging experience), lipid stores, and ovary development, which can facilitate caste-specific collections for future studies. Received 23 May 2008; revised 6 November 2008; accepted 20 November 2008.     

An experimental analysis of pleometrotic advantage in the desert seed-harvester antMessor pergandei (Hymenoptera; Formicidae)

Summary Starting colonies of the desert seed-harvester antMessor pergandei are clumped in the field and face severe intraspecific competition through brood raiding. Single foundress laboratory colonies ofM. pergandei are more likely to succeed at brood raiding with conspecific colonies if they are given additional workers and mature pupae several days prior to brood raiding. Per foundress fecundity remains constant across laboratory starting colonies established with 1, 3 and 5 foundresses. These results suggest that the selective advantage of cooperative colony foundation (pleometrosis) in this and similar species may derive directly from the ability of multiple foundresses to produce a larger brood raiding force.     

Ecological barriers to early colony establishment in three coexisting acacia-ant species in Kenya

In black cotton uplands in East Africa, four symbiotic acacia-ant species compete for possession of a single swollen thorn tree species, Acacia drepanolobium, and yet coexist at fine spatial scales. Three of the four ant species produce independent foundress queens that establish colonies claustrally within swollen thorns, most often on small saplings. We conducted surveys of such saplings at two sites in 2001 and 2004, and examined foundresses and incipient colonies within their swollen thorns to determine what factors influence their success. Competition among foundresses for nest initiation sites was intense, with an average of over one founding attempt per swollen thorn in all samples, and with living and dead queens significantly hyper-dispersed among available thorns. Combat with other foundresses was the most common cause of death among claustral queens, especially for Tetraponera penzigi. In interspecific battles for nest initiation sites, T. penzigi was dominant over Crematogaster nigriceps and C. mimosae, and C. nigriceps won over 80% of its contests with C. mimosae foundresses. For singleton foundresses, brood parasitism by the braconid wasp Trigastrotheca laikipiensis typically results in the death of the entire ant brood. Host queens defend parasite larvae, pupae and eclosed adults, apparently unable to distinguish the wasps from legitimate offspring. Rates of brood parasitism were as high as 15–20% for incipient colonies of both Crematogaster species, but were extremely low for T. nigriceps in all samples. Although T. penzigi and C. nigriceps foundresses are always solitary, approximately 18% of claustral C. mimosae colonies contain cooperating pleometrotic queens. For unparasitized, claustral C. mimosae colonies, brood production per queen did not differ between solitary and cooperating foundresses. However, the per-capita risks associated with parastitism were reduced for pleometrotic queens. Received 8 March 2005; revised 23 May 2005; accepted 3 June 2005.     

Nest site selection by fire ant queens

Summary We examined nest site selection by foundresses of the polygyne form ofSolenopsis invicta. In the laboratory, foundresses were allowed to nest in control soil vs. soil inoculated with refuse from a colony ofPheidole dentata, a natural enemy. In a second experiment foundresses chose between control andS. invicta soil. More foundresses (100%) avoided the soil of a natural enemy than the soil of a conspecific colony (79%) that may accept them into the nest. Foundresses from this polygyne ant appear to avoid other colonies in general, but avoid heterospecific colonies more.     

Induction of colony initiation by Japanese native bumble bees using cocoons of the exotic bumblebee Bombus terrestris

The colony initiation rates of Bombus hypocrita (a native Japanese bumblebee) and Bombus terrestris (a European species) foundresses were compared after 4 weeks of exposure to B. terrestris cocoons. The B. terrestris cocoons, when replaced weekly, were effective for inducing oviposition by foundresses of both species. There were no significant differences in the colony initiation rates of B. terrestris and B. hypocrita, either with the control treatment or with the cocoons. The cocoon method was also tested for five species and two subspecies of native Japanese bumblebees. The colony initiation rate was higher for foundresses of the subgenus Bombus s. str. than for foundresses of the subgenera Pyrobombus, Diversobombus, and Thoracobombus. When replaced weekly, the cocoons of B. terrestris are effective inducers of colony foundation in three Japanese native species, namely B. ignitus, B. hypocrita hypocrita, and B. h. sapporoensis.     

Co‐foundress confinement elicits kinship effects in a naturally sub‐social parasitoid

Kinship among interacting individuals is often associated with sociality and also with sex ratio effects. Parasitoids in the bethylid genus Goniozus are sub‐social, with single foundress females exhibiting post‐ovipositional maternal care via short‐term aggressive host and brood defence against conspecific females. Due to local mate competition (LMC) and broods normally being produced by a single foundress, sex ratios are female‐biased. Contests between adult females are, however, not normally fatal, and aggression is reduced when competing females are kin, raising the possibility of multi‐foundress reproduction on some hosts. Here, we screen for further life‐history effects of kinship by varying the numbers and relatedness of foundresses confined together with a host resource and also by varying the size of host. We confined groups of 1–8 Goniozus nephantidis females together with a host for 5+ days. Multi‐foundress groups were either all siblings or all nonsiblings. Our chief expectations included that competition for resources would be more intense among larger foundress groups but diminished by both larger host size and closer foundress relatedness, affecting both foundress mortality and reproductive output. From classical LMC theory, we expected that offspring group sex ratios would be less female‐biased when there were more foundresses, and from extended LMC theory, we expected that sex ratios would be more female‐biased when foundresses were close kin. We found that confinement led to the death of some females (11% overall) but only when host resources were most limiting. Mortality of foundresses was less common when foundresses were siblings. Developmental mortality among offspring was considerably higher in multi‐foundress clutches but was unaffected by foundress relatedness. Groups of sibling foundresses collectively produced similar numbers of offspring to nonsibling groups. There was little advantage for individual females to reproduce in multi‐foundress groups: single foundresses suppressed even the largest hosts presented and had the highest per capita production of adult offspring. Despite single foundress reproduction being the norm, G. nephantidis females in multi‐foundress groups appear to attune sex allocation according to both foundress number and foundress relatedness: broods produced by sibling foundresses had sex ratios similar to broods produced by single foundresses (ca. 11% males), whereas the sex ratios of broods produced by nonsibling females were approximately 20% higher and broadly increased with foundress number. We conclude that relatedness and host size may combine to reduce selection against communal reproduction on hosts and that, unlike other studied parasitoids, G. nephantidis sex ratios conform to predictions of both classical and extended LMC theories.     

Modes of colony foundation by females of different morphotypes in the paper wasps (Hymenoptera,Vespidae, <Emphasis Type="Italic">Polistes</Emphasis> Latr.)

Paper wasps of the genus Polistes exhibit wide variability both of the color and size of foundresses, and the mode of colony foundation. The colony foundation and social hierarchy were studied in populations of two Palaearctic species, Polistes dominulus (Christ) and P. nimphus (Christ) (April–May 2006, the Crimean Peninsula, Ukraine), and one Neotropical species, P. lanio (F.) (October 2004–January 2005, Trinidad Island). It was shown that foundresses of various color and size morphs tended to found nests in a peculiar way: singly (haplometrosis), by groups (pleometrosis), or occupying several nests simultaneously (polycaly). The relationship between coloration, size, and mode of colony foundation was species-specific. P. dominulus foundresses with a darker clypeus and a paler mesonotum preferred to found nests alone and had a dominant position in pleometrosis. The haplometrotic P. nimphus foundresses more often displayed darker variants of the clypeus and paler variants of the mesonotum than did foundresses from pleometrotic colonies. P. lanio foundresses from pleometrotic colonies differed from those from polycalic colonies in the coloration variability of the scutum, propodeum, and the 2nd metasomal tergite. The dominant and subordinate P. nimphus and P. lanio foundresses differed in the size of head and wings. The possible significance of the subdivision of foundresses into the “generalists” and “specialists” for the variability structuring in the population is considered.     

Lateral Vibrations by Social Wasps Signal Larvae to Withhold Salivary Secretions (Polistes fuscatus, Hymenoptera: Vespidae)

The communicative meaning of an oscillation (lateral vibration) performed by foundresses of social wasps (Polistes fuscatus) was investigated by sampling larval saliva from 12 postworker, single-foundress colonies and by viewing videotapes of 17 multiple-foundress (139.3 h) and 16 single-foundress (32 h) preworker colonies. Foundresses spent significantly more time inspecting cells before performing a lateral vibration (LV) than after and commonly became inactive after performing an LV. The proportion of departures temporally proximate to LVs was significantly greater in single- than in multiple-foundress colonies. Departures and LVs were significantly temporally associated in single-foundress colonies. Single foundresses were significantly more likely to perform an LV prior to departing on longer than shorter trips. In multiple-foundress colonies there was a significant temporal relationship between LVs and departures that left the nest unattended. Larvae secreted significantly less saliva, a nutritious substance provided to adults, immediately after an LV than in the absence of an LV. Contextual evidence and results of sampling larval saliva indicate that LVs by foundresses signal larvae to withhold or reduce their secretion of saliva.     

Costs and benefits of joint colony founding in Australian Acacia thrips

Facultative joint colony founding by social insects (pleometrosis) provides an outstanding opportunity to analyze the costs and benefits of sociality. Pleometrosis has been documented for a range of social insects, but most studies on the adaptive benefits of this behavior are restricted to the Hymenoptera. In this study, we provide the first analysis of costs and benefits associated with pleometrosis for Australian Dunatothrips, which form domiciles by glueing together phyllodes (leaves) of their Acacia host plant. In Dunatothrips aneurae, the distribution of foundress numbers per nest indicated that females formed associations non-randomly. Furthermore, average group size was independent of both the number of foundresses on the host plant and the number of mature colonies, suggesting that this behavior was not simply a response to limited availability of nesting sites. Although per capita reproduction declined with increasing group size, we also identified two benefits of pleometrosis: (1) individual foundresses in groups had higher survival than solitary foundresses during the brood development period, and (2) larger colony sizes resulting from pleometrosis provided a benefit later in colony development, because a higher proportion of D. aneurae adults survived invasions by the kleptoparasite Xaniothrips mulga when colony size was larger. These results demonstrate that the reproductive costs of pleometrosis are at least partially counterbalanced by survival benefits. Received 4 April 2006; revised 9 September 2006; accepted 20 September 2006.     

Cooperative foundation of colonies by unrelated foundresses in the antPolyrhachis moesta

Summary Field studies ofPolyrhachis moesta revealed that colonies were facultatively founded by more than one queen, even though each established colony was inhabited by only one queen with her workers. In pleometrotic colonies, it was found that cofoundresses frequently exhibited cooperative behaviors, including trophallaxis. The results of DNA fingerprinting showed that multiple foundresses were not genetically to one another. It seems that they aggregate randomly and independently of their relatedness. A discussion of the reasons why foundresses ofP. moesta do not aggregate preferentially with genetically related females.     

Nest Hydrocarbons as Cues for Philopatry in a Paper Wasp

Philopatric behavior has been demonstrated in a wide taxonomic spread of animals. In temperate environments, overwintered Polistes wasp foundresses often return to their natal nest prior to initiating colony construction. Previous research has shown that these spring foundresses can identify the natal nest in the absence of landmark and gross morphological cues. Hydrocarbons are essential recognition cues for Polistes nest and nestmate discrimination, but cuticular hydrocarbon profiles can become homogenized when foundresses overwinter in mixed colony groups. We examined the hydrocarbon profiles of Polistes dominulus foundresses and nests before and after an overwintering period, and found that the hydrocarbon profiles of nests remain unique over time and that this uniqueness is influenced by the original foundresses. Our data raise the possibility that in returning to the natal nest, foundresses reacquire their colony‐specific signature, which may play a role in the formation of cooperative associations.     

Rank and epicuticular hydrocarbons in different populations of the paper wasp <Emphasis Type="Italic">Polistes dominulus</Emphasis> (Christ) (Hymenoptera,Vespidae)

Summary Several studies of social insects have shown that epicuticular hydrocarbons are involved in recognition. The hypothesis is that the animals can use differences in chemical composition to acquire information about conspecific status (sex, colony, reproductive status or caste recognition).In this study, we searched for differences between the epicuticular profiles of alpha and beta co-foundresses in Polistes dominulus (Christ) colonies from three separate localities. Our aim was to identify specific compounds or sets of compounds which could be general indexes of foundress rank position in an associative foundation.We found quantitative and qualitative differences in the epicuticular profile among the three populations. The compounds that differentiate between alpha foundresses and their subordinates in the two neighbouring localities were not the same as in the third population. However, in all localities the alpha foundresses of each associative foundation presented a higher proportion of heavy compounds than their respective subordinates. Chemical differences related to rank position may be a fertility signal and are probably caused by physiological and behavioural characteristics, although it is not yet certain if they are actually used by the wasps.Received 28 November 2003; revised 22 February 2004; accepted 25 February 2004.     

Behaviour in usurpers and late joiners ofPolistes biglumis bimaculatus (Hymenoptera,Vespidae)

Summary During the late pre-emergence phase, a foundress of the paper waspPolistes biglumis bimaculatus may be expelled by a conspecific female from her own nest (usurpation) or, less frequently, joined by another female of the same species (late association). The behaviour of femalePolistes biglumis bimaculatus, when usurping a conspecific colony or joining another foundress, is compared with that of foundresses on non-usurped colonies. The most conspicous difference is the intense abdomen stroking behaviour the usurper performs over the comb surface on the first days after usurpation. As observed in otherPolistes species, once usurpers and joiners arrive on a strange nest they will destroy most of the immature brood of the previous nest owner. Although host workers are not aggressive towards the intruder females, reproductive success of usurpers and joiners is low compared with that of legitimate foundresses. The same behaviours observed on usurped colonies are found in the obligate social parasites ofPolistes. These behaviours are therefore discussed in the context of the evolution of intra- and inter-specific parasitism.     

The triple alliance: how a plant-ant, living in an ant-plant, acquires the third partner, a scale insect

Summary. Mutualistic associations between ants and plants often involve a third partner, scale insects (Hemiptera, Coccoidea). In southeast Asia, plant-ants of the genus Cladomyrma live together with coccoids in hollowed twigs of a wide range of ant-plants (myrmecophytes). Established colonies never lack sap-sucking scale insects and the ants appear to be dependent on the honeydew excretions of their trophobionts. Acquisition of scale insects thus seems to be an important step in the successful establishment of a new colony on a host plant. Coccoids may either be transported by a foundress ant on her nuptial flight (vertical transmission via co-dispersal) or ant and coccoid disperse on their own and associate anew during colony initiation (horizontal transmission). Here we test the assumption that founding gynes of Cladomyrma do not transport scale insects during their dispersal flight, and we determine when and how trophobionts are acquired. The results obtained in this study show that although coccoids are not carried by foundresses, their numbers within founding chambers of Cladomyrma increased with the founding stage. In all Cladomyrma species, colony-founding is claustral and the entrance hole into the nest chamber is first plugged with pith debris and later grows over by wound callus. However, access of trophobionts into hollow twigs is facilitated either by additional small openings (secondary slits) produced by the foundress ant after hollowing out a nest chamber or by the maintenance of an orifice at the entrance hole during the founding period. Immature pseudococcids experimentally placed onto twigs close to a Cladomyrma foundation readily entered these slits. Exclusion experiments suggest that foundresses of C. petalae are less successful in rearing their first offspring if entry of coccoids into founding chambers is prevented.Received 28 September 2003; revised 9 May and 11 October 2004; accepted 9 November 2004.     

Consequences of foundress aggregation in the brown citrus aphid Toxoptera citricida

1. A field study was conducted in which colonies of Toxoptera citricida (Kirkaldy) were initiated with different numbers of foundresses on ungrafted citrus trees var. ‘Carrizo’ within an existing citrus grove in central Florida. 2. The growth rate, longevity, and final size of T. citricida colonies were all positively correlated with number of foundresses. 3. Colonies initiated by many foundresses produced alatae earlier than those initiated by fewer foundresses. 4. Nymphal developmental time was reduced in colonies initiated by more foundresses but the mean number of grand‐progeny per foundress was lower when only surviving colonies were compared. 5. The longevity of foundresses was unaffected by their number but foundress fecundity and nymphal survival decreased as foundress number increased. 6. The most abundant natural enemies were Diptera: Syrphidae, primarily Pseudodorus clavatus (F.), and Chamaemiidae (Leucopis sp.), followed by Coleoptera: Coccinellidae, primarily Cycloneda sanguinea (L). 7. Colonies initiated with more foundresses attracted more oviposition by dipteran predators than did colonies initiated with fewer foundresses.     

茉莉酸甲酯诱导大戟三萜类代谢的研究

京大戟是多年生草本药用植物,入药部分是其干燥根,但可入药的京大戟资源由于生长缓慢以及环境污染的加剧而越发匮乏,因此解决大戟资源日益紧张的问题是当今药用植物资源开发与利用方向的重要课题。京大戟含有三萜类、二萜类、黄酮类等丰富的活性成分,一些常见药用植物的有效成分是三萜类化合物,其在抗病毒、抗肿瘤、免疫调节等方面具有很好的活性。对植物萜类物质代谢起重要作用的关键酶,如3-羟基,3-甲基戊二酰辅酶A还原酶(hmgr)、鲨烯合酶(sqs)、法尼基焦磷酸合酶(fps)的基因克隆及活性研究取得了进展和突破,但通过调控萜类物质代谢途径中关键酶基因的表达来诱导终产物合成的研究鲜有报道。通过研究大戟萜类物质代谢途径进而利用基因工程手段提升目的物质的产量来解决京大戟药源短缺问题具有重要意义。该研究以大戟愈伤组织为材料,使用茉莉酸甲酯分别按时间梯度和浓度梯度进行诱导,将诱导后的愈伤组织分为两部分:一部分提取其总RNA,以actin为内参基因进行反转录,实时定量RT-PCR分析大戟三萜类代谢途径中hmgr、sqs与fps基因的相对表达差异;另一部分用于提取其总三萜并使用分光光度法进行含量测定。实时定量RT-PCR分析结果表明,茉莉酸甲酯可诱导3个基因的表达,但其表达模式不一样。相应的京大戟愈伤组织中总三萜的含量明显提高,最高可较未处理样品增加27%。研究结果可为茉莉酸甲酯促进药用植物大戟三萜类物质积累的分子机制研究提供参考。     

Fungus garden platforms improve hygiene during nest establishment in <Emphasis Type="Italic">Acromyrmex</Emphasis> ants (Hymenoptera,Formicidae, Attini)

Physically isolating organisms from disease agents should reduce the likelihood of disease transmission and infection, and increase survival and growth, particularly in more vulnerable, early ontogenetic stages. During nest founding in fungus-growing ants, foundresses of most genera use a garden platform to isolate the incipient fungal garden from the soil of the underground chamber. We studied nest founding in Acromyrmex octospinosus to test the hypotheses that the use of a platform (rootlets used to suspend the fungus garden): (i) reduces the likelihood that the garden will be contaminated by soil-borne microbial pathogens; (ii) results in more rapid growth of a young colony; and (iii) increases colony survival. We manipulated natural incipient nests to have gardens either in contact with or isolated from soil surrounding the chamber, and nests with and without foundresses present. We found a higher incidence of infection in gardens that were in contact with the chamber soil and without queens, compared with gardens isolated from the chamber soil with and without foundresses. The production of eggs, larvae and pupae, as well as leaf area harvested, were significantly different between nests with and without platforms, but there were no differences in the production of workers nor garden biomass. Likewise, there were no differences between treatments in colony survival rates over 8 weeks. Using smaller incipient gardens, however, gardens with and without platforms differed in survivorship rates after 24 hours. The results indicate that the use of a platform to cultivate an incipient fungal garden is an adaptation to reduce soil-borne diseases and increase colony performance. Received 28 July 2006; revised 15 November 2006; accepted 22 November 2006.