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

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

Co-evolution-driven cuticular hydrocarbon variation between the slave-making ant Rossomyrmex minuchae and its host Proformica longiseta (Hymenoptera: Formicidae)

Summary. Proformica longiseta exists as two populations in the Sierra Nevada Mountains in Spain, only one of which is parasitized by the slave-maker ant Rossomyrmex minuchae. To investigate the possible effect of co-evolutionary pressures on cuticular hydrocarbon (CHC) profiles (the presumed nestmate recognition cues), we performed a comparative analysis of the CHC of R. minuchae and P. longiseta colonies from both the allopatric and sympatric populations; the latter includes samples of enslaved as well as free-living workers. Discriminant analyses based on these chemical profiles showed two clear profile groups: the first comprised R. minuchae and both enslaved and free-living P. longiseta from the sympatric population; and the second the allopatric P. longiseta workers. As expected, the profiles of the two sympatric P. longiseta groups (enslaved and free-living) were distinct; but, interestingly, those of the enslaved P. longiseta and its parasite R. minuchae were also distinguishable. This indicates that despite their cohabitation each species maintains its own chemical identity. Profile similarity between the sympatric free-living P. longiseta and its parasite may explain the lower than expected aggression observed during raids. We further speculate that in view of the differences between the sympatric and allopatric population of P. longiseta, co-evolutionary pressures have driven changes in the profile of the former to better match that of its parasite R. minuchae. Such an adjustment may have enabled nests of the sympatric P. longiseta to endure multiple raids by the parasite (due to the reduced aggression) and thus to continue to reproduce despite the damage inflicted by the raids.     

Information transfer during recruitment in the ant<Emphasis Type="Italic"> Lasius niger</Emphasis> L. (Hymenoptera: Formicidae)

In this paper, we used the food-correlated search behavior observed in foraging ants returning to a previously rewarding site to study information transfer during recruitment in the ant Lasius niger. We hypothesized that, if information about the characteristics of the food is conveyed during recruitment, food-correlated search tactics should also be observed in recruited workers. Our results show that the characteristics of the trajectories of recruited workers are comparable to those of scout ants returning to a site or prior food find and depend more on the type (prey/sugar) than on the quality (sugar concentration) of the food discovered by the scouts. Independent of sugar concentration, workers recruited to a source of sugar search with a greater sinuosity than workers recruited to a prey. Experimental manipulation of the recruitment signals (chemical trail and contact between ants) shows that the trail pheromone laid down by recruiting ants does not play a role in the modification of trajectory sinuosity. This change appears to be most likely triggered by a direct perception of the residue of sugar smeared on the body of the recruiting workers coming back to the nest.Communicated by J. Heinze     

Chemical camouflage by myrmecophilous beetles Zyras comes (Coleoptera: Staphylinidae) and Diaritiger fossulatus (Coleoptera: Pselaphidae) to be integrated into the nest of Lasius fuliginosus (Hymenoptera: Formicidae)

Summary. The myrmecophilous beetles, Zyras comes (Staphylinidae) and Diaritiger fossulatus (Pselaphidae) are guests of the black shining ant Lasius fuliginosus. Host worker ants never attacked these beetles, and often gave regurgitant to Z. comes following tactile communication with the beetle. By contrast, the workers from colonies without the myrmecophiles showed hostile responses towards Z. comes before tactile contact, but were not aware of D. fossulatus until contact. In L. fuliginosus, workers within a colony shared profiles, but the profiles differed among colonies. GC analyses showed that both Z. comes and D. fossulatus beetles had the same hydrocarbons as L. fuliginosus, and the profiles were more similar to those of the host colony workers than the foreign workers. Both Z. comes and D. fossulatus appear to imitate the hydrocarbon profile of their host workers, allowing integrating into the host nest. A Y-maze bioassay indicated that Z. comes can follow the trail pheromone of L. fuliginosus. This suggests that Z. comes may detect other chemical signals of L. fuliginosus to keep closer interactions with the workers. Received 22 June 2001; accepted 12 November 2001.     

Species-specificity in trail pheromones and Dufour's gland contents ofCamponotus atriceps andC. floridanus (Hymenoptera: Formicidae)

Summary By means of gas chromatography, gas chromatographic coupled mass spectrometry, trail-following experiments and electrophysiological recordings from worker antennae, the major trail pheromone components from the hindgut of the formicine speciesCamponotus atriceps andC. floridanus were identified as 3,5-dimethyl-6-(1-methylpropyl)-tetrahydropyran-2-one and nerolic acid, respectively. The Dufour's gland contents of both species, investigated by gas chromatographic coupled mass spectrometry, show significant differences.Pheromones 104: Janssen E, Übler E, Bauriegel L, Kern F, Bestmann H-J, Attygalle AB, Steghaus-Kovacs S, Maschwitz U: Trail pheromone of the Ponerinae antLeptogenys peuqueti (Hymenoptera: Formicidae): a multicomponent mixture of related compounds     

Chemical mimicry in the root aphid parasitoid Paralipsis eikoae Yasumatsu (Hymenoptera: Aphidiidae) of the aphid-attending ant Lasius sakagamii Yamauchi & Hayashida (Hymenoptera: Formicidae)

Summary. An aphidiid wasp, Paralipsis eikoae, was associated with both Lasius niger and L. sakagamii attending the wormwood root aphid Sappaphis piri. An L. sakagamii worker was observed carrying a winged female P. eikoae to its nest with its mandible, but it did not kill the wasp. Once accepted by the ants, the wasp often mounted and rubbed against the worker ants and sometimes teased them to regurgitate food to itself. No workers in the colony attacked the wasp. Conspecific foreign workers, however, viciously attacked the wasp when encountered. Gas chromatography-mass spectrometry analyses showed that the accepted wasp had complex cuticular hydrocarbons that were very similar to those of its host ants, whereas the winged wasps collected outside the ant nest showed only a series of n-alkanes. Additionally, the accepted wasp had a hydrocarbon profile closer to that of its host ants than to the conspecific foreign ants. We believe the wasp mimics ant cuticular hydrocarbons to integrate into the ant nest, acquiring the hydrocarbons by mounting and rubbing against the ants. In contrast, the cuticular hydrocarbons of the emerged wasp contained larval and pupal hydrocarbons of L. sakagamii that were also similar to those of L. niger. Both ant species rejected adult workers of the other species but accepted their larvae and pupae. We suggest that the emerged P. eikoae mimics the cuticular hydrocarbons of these Lasius larvae and pupae, which allows P. eikoae to be accepted by both L. sakagamii and L. niger. Received 11 March 1998; accepted 22 July 1998.     

Chemistry of the postpharyngeal gland secretion and its implication for the phylogeny of IberianCataglyphis species (Hymenoptera: Formicidae)

Summary A comparative morphological and chemical study of six endemic species ofCataglyphis from the Iberian Peninsula:C. ibericus, C. rosenhaueri, C. hispanicus, C. humeya, C. velox andC. floricola and the Moroccan speciesC. bombycinus is described. The morphological study relied primarily on genitalia characteristics, whereas the chemical study concentrated on the postpharyngeal gland constituents. Cladograms based on the morphological and chemical data were performed using Ward's method. The dendrogram based on morphological features revealed that the IberianCataglyphis can be classified into three species groupsalbicans, altisquamis andemmae. The same pattern occurred when the dimethylalkanes constituents of the postpharyngeal gland were utilised as character states, with a slight displacement of species within thealtisquamis group. However, when the complete hydrocarbon blend was utilised major discrepancies in the dendrograms occurred.Cataglyphis velox proved to be very similar toC. bombycinus, whereasC. floricola clustered with the other two species of thealtisquamis group. Based on the geographical distribution and paleontological data (Tinaut 1993) it is assumed thatC. floricola recently invaded the Iberian Peninsula. Based on the chemical findings we postulate that chemical character displacement occurred inC. floricola as a result of its sympatry withC. velox after the former colonized the Iberian Peninsula. We further discuss the possible reason for the different dendrograms obtained when only the dimethylalkanes are considered and its implication for the communicative role of the postpharyngeal gland secretion in these ants.     

Predatory behavior and chemical communication in two Metapone species (Hymenoptera:Formicidae)

Summary. Colonies of two species of Metapone (M. madagascarica, M. new species.) were collected in Madagascar and established in laboratory nests. It could be demonstrated that both species are specialist predators of termites (Cryptotermes kirbyi). During hunting the ants sting the termites and thereby paralyze and preserve the prey alive. In this way prey can be stored in the ant nest for extended periods. During foraging and colony emigrations the ants lay chemical trails with poison gland secretions. Among the seven compounds identified in the venom only methyl pyrrole-2-carboxylate elicits trail following behavior in both Metapone species. Received 11 February 2002, accepted 23 February 2002.     

Complex chemical communication in the crazy ant <Emphasis Type="Italic">Paratrechina longicornis</Emphasis> Latreille (Hymenoptera: Formicidae)

Summary. The formicine ant Paratrechina longicornis is known for its extremely opportunistic foraging behaviour. Only a single trail pheromone source, the rectum, was previously described from this ant. Our detailed examination of this ant’s chemical communication system revealed the presence of at least four sources of pheromones. Rectum, poison sac, and Dufour gland contain orientation components with decreasing effectiveness and persistence (in the sequence mentioned) as well as attractants with increasing effectiveness. Furthermore, the mandibular gland contains repellents, and a releaser of defensive behaviour. This set of various signals of different strength and persistence allows an extraordinary degree of flexibility and efficiency in the collective behaviour of P. longicornis, especially food exploitation, and thus may contribute to this insect’s overall ecological success.     

Profiles of cuticular hydrocarbons mediate male mate choice and sexual isolation between hybridising <Emphasis Type="Italic">Chrysochus</Emphasis> (Coleoptera: Chrysomelidae)

Summary. Chemical signals frequently underlie sexual isolation between insect species. Our understanding of the evolutionary forces influencing these signaling systems is known for very few systems, challenging both our efforts to understand insect speciation, and our ability to predict long-term changes in the chemical communication systems of insects. Thus, we are in need of more systems in which both the chemical signals causing sexual isolation and the evolutionary forces driving sexual isolation are understood. Sexual isolation in the hybrid zone between Chrysochus cobaltinus and C. auratus has apparently increased in response to natural selection against hybridisation (i.e. reinforcement). Previous experiments suggested that this isolation was due, at least in part, to male preferences for conspecific females. Here, we confirm this role of male choice, and document that male mate choice in this system is influenced by cuticular hydrocarbon (CHC) profiles. Specifically, male C. cobaltinus responses to control cadavers and conspecific female cadavers painted with different cuticular hexane extracts, together with analyses of the composition of those extracts, revealed that male mate choice is governed by CHC profiles. Multivariate analyses of GC profiles demonstrated that those profiles are indeed both sex- and species-specific. Although GC-MS enabled identification and quantification of the specific cuticular hydrocarbons, we have not yet determined which individual compounds govern mate choice. Having established that CHCs influence sexual isolation in this system, we can now assess the evolutionary lability of these cues, which will inform both our understanding of speciation, and of the conditions under which the chemical signaling systems that influence mate choice in insects can evolve.     

Chemical mimicry of cuticular hydrocarbons – how does Eremostibes opacus gain access to breeding burrows of its host Parastizopus armaticeps (Coleoptera, Tenebrionidae)?

Summary. Breeding burrows of Parastizopus armaticeps armaticeps, a fossorial desert tenebrionid beetle, are cleptoparasitised by the closely related Eremostibes opacus. Gas chromatographic analyses show a high congruity of the cuticular hydrocarbons of both species. We compare these hydrocarbon patterns with those of four other Stizopina species and the Scaurini Herpiscius sommeri. In a bioassay, dummies treated with cuticular hydrocarbon extracts of E. opacus and the P. a. bifidus parasite E. bushmanicus were mostly ignored by P. a. armaticeps, whereas dummies with applied extracts of the remaining species were heavily attacked. We show that there is a correlation between agonistic behaviour of P. a. armaticeps towards the intruder and the chemical similarity of the cuticular hydrocarbons of the two species. Furthermore, we produced quantitatively modified hydrocarbon patterns of E. barbatus by changing the temperature at which this species was kept. The new 30 °C type was chemically similar to E. opacus, and was frequently ignored by P. a. armaticeps, whereas a reduction of the temperature to 20 °C only had minor effects on the hydrocarbon pattern. Furthermore, we show that the addition of one single component, heptacosane, to the cuticular hydrocarbon extract of E. opacus alters the host’s reaction. We discuss the role of cuticular hydrocarbons for the recognition of this host-parasite system and the relevance of quantitative characters in the hydrocarbon pattern for the discrimination of the host.     

Do these eggs smell funny to you?: an experimental study of egg discrimination by hosts of the social parasite Polyergus breviceps (Hymenoptera: Formicidae)

Social parasites exploit the behaviours of other social species. Infiltration of host systems involves a variety of mechanisms depending on the conditions within the host society and the needs of the social parasite. For many species of socially parasitic ants, colony establishment entails the usurpation of colonies of other species. This frequently involves the eviction or death of the host colony queen and the subsequent adoption of the invading queen. The social parasite queen achieves host worker acceptance by either manipulating the nest-mate recognition processes of the host or undergoing chemical modification. Little is known, however, about how host workers respond to social parasite eggs or whether host species defend against brood parasitism during parasite invasions. Host species are believed to adopt social parasite offspring because the recent common ancestry between many social parasites and their hosts may grant the sharing of certain characteristics such as chemical cues. Use of multiple host species, however, suggests other processes are needed for the social bond between host and parasite young to form. This study reports the findings of adoption bioassays in which eggs from a slave-maker ant, Polyergus breviceps, were offered to workers of two of its host species from unparasitised or newly parasitised nests to determine whether P. breviceps eggs generally elicit rearing behaviours from multiple host species. Comparisons of parasite egg survival until adulthood with conspecific egg survival reveal that workers of both host species, free-living or newly enslaved, do not typically accept slave-maker eggs. Both host species thus have sufficient discriminatory power to reject social parasite eggs although our hydrocarbon analysis indicates parasite eggs may be adapted to their local host species. Combined these results suggest that host rearing of P. breviceps eggs may reflect an evolutionary equilibrium that is maintained by probability and cost of recognition errors.Communicated by L. Sundström     

Queen dimorphism and reproductive strategies in the fire ant Solenopsis geminata (Hymenoptera: Formicidae)

Alate trapping studies of a monogyne population of the fire ant Solenopsis geminata indicate that two sizes of gynes are produced. Macrogynes, which participate in late spring and summer mating flights, are larger, fattier, and more than twice as heavy as microgynes, which participate in fall mating flights. Three patterns of gyne production were observed in 51 colonies studied: 35 produced macrogynes only, 9 produced microgynes only, and 7 produced both morphs, contributing to both summer and fall mating flights. Behavioral evidence and rearing studies suggest that macrogynes found new colonies independently, whereas microgynes achieve colony queen status by infiltrating or being adopted by established colonies. Of the total number of female alates collected from the trapped colonies, 56% were microgynes. However, because of their smaller size and lower fat content, microgynes made up only one-third of the caloric investment in female alates. By measuring the thorax lengths of queens from mature colonies, we determined that at least 56% were macrogynes and 35% or more were microgynes. These results indicate that as a reproductive strategy, colony investment in microgyne production may have at least as high a payoff as investment in macrogyne production.This is publication #24 of the Fire Ant Research Team     

Tarsal secretion marks food sources in bumblebees (Hymenoptera: Apidae)

Summary Scent-marking of artificial food sources by workers of the bumblebee,Bombus terrestris, was investigated. Odour marks deposited on artificial flowers were subsequently collected and chemically analysed. Alkanes and alkenes were identified as the main components. The behaviour mediating capacity of synthetic mixtures of the identified compounds was bioassayed using an artificial flower system. A mixture of alkanes and alkenes, close to natural proportions, released regular foraging behaviour. The reaction proved to be dose-dependent.     

Combined use of pheromone trails and visual landmarks by the common garden ant Lasius niger

This study investigated the relative importance of pheromone trails and visual landmarks on the ability of Lasius niger foragers to relocate a previously used food source. Colonies formed foraging trails to a 1-M sucrose feeder. Sections of this trail were then presented back to the same colony after variable time intervals. Individual outgoing foragers were observed to determine if they walked for 15 cm in the direction of the feeder or not. On newly established pheromone trails formed by 500 ant passages, 77% of the foragers walked in the correct direction vs 31% for control foragers (no trail pheromone). Pheromone trails decayed to the control levels in 20–24 h. Trails formed with fewer ant passages (125 or 30) decayed quicker. The use of visual landmarks was investigated by using trails with outgoing foragers from the colony that established the trail, either in the same room or in a different room, with different visual landmarks, to that used during trail establishment. Approximately 20% more ants walked in the correct direction in the same room vs the different room. This difference decreased to around 10% 2 h after trail establishment, indicating that the ants in the different room were learning the new visual cues to navigate by. Our results show that visual landmarks and pheromone trails are approximately equally useful in initially guiding L. niger foragers to food locations and that these two information sources have a complementary function.     

Heritable pheromone blend variation in a local population of the butterbur borer moth Ostrinia zaguliaevi (Lepidoptera: Crambidae)

Summary. In a local population of Ostrinia zaguliaevi Mutuura & Munroe (Lepidoptera: Crambidae), extensive variation was found in the blend ratio of three sex pheromone components, (Z)-9-tetradecenyl acetate (Z9-14:OAc, 10.2-63.8%), (Z)-11-tetradecenyl acetate (Z11-14:OAc, 32.2-86.8%), and (E)-11-tetradecenyl acetate (E11-14:OAc, 2.1-11.9%). The variation was observed over a three-year period (2002-2004). Mother-daughter regression analyses have shown that although the heritability of the minor component E11-14:OAc was not significant, the heritability of the proportions of Z9-14:OAc and Z11-14:OAc were substantial (0.5–0.6). In artificial selection experiments, the mean % Z9-14:OAc in the sex pheromone changed significantly within three generations (37% in the control line, 48 and 52% for two lines selected for increase, and 23 and 30% for two lines selected for decrease). Despite these changes, the amounts of fatty acyl pheromone precursors, (Z)-9-, (Z)-11- and (E)-11-tetradecenoate, in the pheromone gland were not significantly affected by the selection. Taken together, variation in the pheromone blend of O. zaguliaevi is likely to be attributable to a few genes involved in the reduction or acetylation of fatty acyl pheromone precursors, the last two steps in pheromone biosynthesis.     

Toxicity of the venom in three neotropical Crematogaster ants (Formicidae: Myrmicinae)

Bioassays were performed to investigate and compare the toxic properties of the contact venoms of three Neotropical Crematogaster species (C. sp. prox. abstinens, C. distans and C. brevispinosa rochai), whose main venom compounds are chemically different (long chain derivatives linked to an aldehyde or a primary acetate, and furanocembrenoid diterpenes, respectively). Different quantities of venom were topically applied on the bodies of three target insect species (workers of the termite Nasutitermes ephratae, workers of the ant Solenopsis sp., “media” and “major” workers of the ant Cephalotes pusillus). The toxicity of the venom greatly varied according to the Crematogaster species, the venom of C. b. rochai showing the highest toxic activity towards all target species. The sensitivity of the target species also greatly varied, the workers of N. ephratae showing a great sensitivity to the venoms, while the heavily armoured workers of the ant C. pusillus, particularly “major” ones, were resistant. Cuticle thickness was found to be a key factor in the resistance to the toxic effects of Crematogaster venom. Sensitivity of the target workers was also directly related to the amount of venom applied to their cuticle. Workers of C. distans were resistant to topical application of their own venom, while C. sp. prox. abstinens and C. b. rochai were not. In all cases, the venoms were slow acting, as several hours to one or even several days were needed to obtain a significant mortality. The main biological activities of the Crematogaster venom could be related to more immediate repellent properties, as recently shown with the European species C. scutellaris.     

Hibernation clustering and its consequences for associative nest foundation in<Emphasis Type="Italic"> Polistes dominulus</Emphasis> (Hymenoptera Vespidae)

Various studies indicate that cuticular hydrocarbons (CHCs) act as cues for nestmate recognition in paper wasps and other social insects. A recent study showed that associative nest foundation in Polistes dominulus is mainly performed by foundresses coming from the same locality. In the present study, we induced future foundresses of P. dominulus collected in two different localities to hibernate in the laboratory in aggregates of individuals from the same or different localities. After 2 months of hibernation, foundresses from different localities but from the same experimental cluster did not show any preference to associate, at the time of nest foundation, with individuals coming from the same original locality. The cuticular chemical profiles of individuals from the mixed hibernation clusters were quite similar and significantly different from those of individuals which hibernated with other foundresses from the same locality. These findings suggest that, in this species, mechanisms other than nestmate chemical recognition play a major role in the spring association of gynes during the foundation of a new nest.Communicated by R.F.A. Moritz     

Chemical recognition of queen cells by honey bee workersApis mellifera (Hymenoptera: Apidae)

Summary Honey bee workers are able to nurse or to destroy and thus to recognize the capped queen cells containing a pupa. Fatty acid esters, especially methyl oleate, methyl palmitate and ethyl oleate were found in significant amounts on the queen pupal cuticle. Methyl oleate, the major component, along with smaller amounts of methyl linoleate and methyl linolenate, were involved in the recognition of queen cells by workers. In natural conditions of the colony, queen cells containing a paraffin pupal lure with methyl oleate were accepted 5.9 days by workers, releasing about 1.8 queen pupa equivalents during that period, when control cells (without ester) were kept only 2.1 days. Although these esters are non specific to honey bees, they are of great importance in social regulation of the honey bee colony.     

12种杀菌剂制剂对半闭弯尾姬蜂的急性毒性及安全性评价

采用试管药膜法测定了12种杀菌剂制剂对半闭弯尾姬蜂成蜂的急性接触毒性,结合安全性系数评价了供试药剂对半闭弯尾姬蜂的安全性。急性毒性测定结果表明,氟硅唑和啶菌噁唑对半闭弯尾姬蜂成蜂的接触毒性最高,LC50值分别为220.022和223.115 mg·L-1;其次为多抗霉素、丙森锌和戊唑醇,LC50值分别为436.496、472.358和638.638 mg·L-1;其余7种杀菌剂制剂嘧霉胺、啶酰菌胺、苯醚甲环唑、多菌灵、嘧菌酯、嘧菌环胺和异菌脲对半闭弯尾姬蜂成蜂的触杀毒性都较低,LC50值均大于1 000 mg·L-1。安全性评价结果表明,丙森锌和啶菌噁唑对半闭弯尾姬蜂成蜂具有高风险性,安全系数分别为0.16和0.42;嘧霉胺、多抗霉素、氟硅唑和戊唑醇为中等风险性,安全系数分别为1.43、1.56、2.20和3.23;其余6种杀菌剂制剂啶酰菌胺、苯醚甲环唑、多菌灵、嘧菌酯、嘧菌环胺和异菌脲对半闭弯尾姬蜂表现为低风险性,安全系数均大于5。结果显示:甾醇脱甲基抑制剂氟硅唑、戊唑醇和啶菌噁唑以及苯胺基嘧啶类杀菌剂嘧霉胺、有机硫杀菌剂丙森锌和抗菌素多抗霉素对半闭弯尾姬蜂成蜂具有急性毒性风险,在有害生物综合治理中应谨慎使用,特别是啶菌噁唑和丙森锌,以免对半闭弯尾姬蜂造成不良影响和危害。