Arboreal search in ants: Search on branches (Hymenoptera: Formicidae)

Ants (Formicidae) perform two distinct search behaviors for resources: on the ground they use irregular, almost random alternating looping, and on branches and leaves they resort to outline-tracing (arboreal systematic search), whereby the individual systematically turns to one side at bifurcations and to the opposite side when turning about at end points. Experiments with searching Formica pallidefulva and Crematogaster cerasion artificial stick mazes under seminatural conditions demonstrated that bifurcations and end points only trigger turn decisions, whereas an intrinsic mechanism specifies the handedness of such turns. Arboreal homing differs from arboreal searching by a much stronger tendency to rectify paths by counterturning. The theory is advanced that searching on branches by outline-tracing is evolutionarily derived from ranging search by superposing a sustained intrinsic turn bias and by suppressing random turns.     

Respective contributions of leader and trail during recruitment to food inTetramorium bicarinatum (Hymenoptera: Formicidae)

Summary This paper describes the food recruitment strategy of the antTetramorium bicarinatum, at both the individual and collective levels. The general organisation of recruitment used by this species during the exploitation of sucrose solutions shows similarities with group recruitment described for other species. However, our experiments demonstrate that inT. bicarinatum, the first trail laid by a recruiter during its return trip to the nest is more efficient than in other species using group recruitment, for exampleTetramorium impurum. Moreover, the efficiency of the first trail of aT. bicarinatum recruiter is comparable with that ofTapinoma erraticum, a species that uses mass recruitment. Despite the efficiency of the trail, choice experiments show that the recruited workers prefer to follow the leader rather than the first trail, suggesting the emission of a more attractive signal by the leader on its way back to the food. The function of the leader in this strategy is discussed in terms of collective decisions.     

Arrestment responses ofEretmocerus species andEncarsia deserti (Hymenoptera: Aphelinidae) toBemisia tabaci honeydew

The responses of Eretmocerussp. and Encarsia deserti(Gerling & Rivnay), parasitoids of Bemisia tabaci(Gennadius) to host secretions were investigated. Upon contact with honeydew excreted from the host, the parasitoids' walking speed decreased, whereas the rate of angular turning increased (the walking speed of Eretmocerussp. and Encarsia desertidecreased from 2.0 to 0.8 and from 0.8 to 0.3 mm/s, respectively, and the respective angular turning rates increased from 123.3 to 489.6 and from 267.9 to 979.2 deg/cm). Moreover, the wasps generally turned back toward the secretion as soon as they lost contact with it. This induced arrestment responses resulted in increased searching time by the parasitoids. Laboratory bioassays were conducted in which paper disks in petri dishes containing different amounts of honeydew, equivalent to the secretions of 1, 5, or 50 larvae, were offered to female wasps. The wasps responded by arrestment even to honeydew produced by a single larva. While the response of Eretmocerussp. was quantity dependent, being more intense at higher amounts of honeydew, that of E. desertiwas of equal intensity at all honeydew quantities assayed.     

Oral trophallaxis in adult leaf-cutting ants Acromyrmex subterraneus subterraneus (Hymenoptera, Formicidae)

Adult leaf-cutting ants of the subspecies Acromyrmex subterraneus subterraneus were fed with an Evans Blue dye solution, which allowed the investigation of subsequent exchange of liquids between ants by oral trophallaxis. Trophallactic behavior was filmed and the antennation patterns of donor and recipient ants were described. The ants’ crop capacity was measured following ad libitum feeding on dye solution. Ants previously fed on the dye solution (donors) were placed individually with unfed ants of the same caste (recipients) and the amount of dye solution passed from the donor to the recipient by oral trophallaxis was measured after one hour. The volume received was 0.26 ± 0.15μL (mean ± SD) and the residual volume of dye in the crop of the donors was 0.49 ± 0.23μL. There were 38 trophallactic events recorded for 50 pairs of ants. Trophallaxis was observed from 2.3 min to 21.5 min after initial exposure, with a mean latency of 8.4 ± 5.6 min. The mean duration of a trophallatic event was 2.3 ± 1.3 min. As far as we know, this is the first time that trophallaxis in leaf-cutting ants has been described and quantified. Received 2 December 2005; revised 6 April 2006; accepted 10 April 2006.     

Queen size dimorphism in the ant Tetramorium moravicum (Hymenoptera, Formicidae): Morphometric, molecular genetic and experimental evidence

Summary. By combining different methods we evaluate whether the ant Tetramorium rhenanum is specifically separated from T. moravicum or whether it is a conspecific microgyne form. High-precision morphometry shows a clear difference in queen size. Sequence comparison of 1031 bp of COI reveals that T. rhenanum falls into a clade with T. moravicum, which is significantly separated from T. forte and T. chefketi. T. rhenanum shares at least two haplotypes with T. moravicum and is considered as a junior synonym. Sexual behaviour and colony foundation experiments corroborate conspecificity. The queen dimorphism is discussed in the context of social parameters such as queen number and reproductive strategy.Received 10 June 2004; revised 28 October 2004; accepted 12 November 2004.     

Behavioral response ofGraminella nigrifrons (Homoptera: Cicadellidae) to experimentally manipulated vibrational signals

Mate recognition for the leafhopper Graminella nigrifrons(Forbes) occurs when a male spontaneously emits a multisectional vibrational calling song to which females respond by emitting simple pulses. Significant differences were found among males in the duration, number of chirps, and chirp rate within sections of the song and the total song. Repeatability (proportion of total variation due to differences among males) of call features ranged from very low (0.04 for total chirps in song) to high (0.67 for section 3 chirp rate). However, song modification and playback experiments revealed that the variation in the measured song features was not important in determining whether a female will respond. Rather, female response depended only on the presence of two of the three types of pulses which comprise a chirp. These essential pulses were found within chirps of all call sections that contain chirps. Manipulation of chirp rates from 0.58 to 2.70 times the normal rate did not affect female response, nor did changing the period of silence between the essential pulse types from 0.25 to 1.75 times the normal period. These results suggest that components of the male calling song function in mate recognition but are not used by females to discriminate among conspecific males.     

Effects of Argentine ants (Hymenoptera: Formicidae) on myrmecophilous lycaenid butterfly,Narathura bazalus (Lepidoptera: Lycaenidae), in western Japan

Effects of the invasive Argentine ant, Linepithema humile (Mayr), on a myrmecophilous butterfly, Narathura bazalus (Hewiston), were investigated in the field in western Japan. Larvae of N. bazalus were attended by workers of Argentine ants in invaded parks and of ten native and one cosmopolitan ant species in uninvaded parks. The abundance of eggs and larvae were not significantly different between invaded and uninvaded parks. Pupal weight and parasitized ratio by tachinid flies were also not different between the two types of parks. These results indicate that the role of Argentine ants for the butterfly might be almost equivalent to the native ants.     

A preference assay for quantifying symbiont choice in fungus-growing ants (Attini, Formicidae)

We describe a bioassay for the quantification of cultivar preference (symbiont choice) of fungus-growing ants. The bioassay simultaneously presents mycelium of multiple pure cultivar genotypes to worker ants in a cafeteria-style test arena, and preferred versus non-preferred cultivar genotypes can then be identified based on the ants’ quantifiable behavioral tendencies to convert any of the offered mycelium into a fungus garden. Under natural conditions, fungus-growing ants are likely to express such cultivar preferences when mutant cultivars arise in a garden, or when colonies acquire a novel cultivar from a neighboring colony to replace their resident cultivar. We show that workers from different nests of the fungus-growing ant Cyphomyrmex costatus exhibit repeatable preferences vis-à-vis specific cultivar genotypes. The identified preferred and rejected cultivars can then be used in a performance assay to test whether the ants prefer cultivar genotypes that are superior in enhancing colony fitness (measured, for example, as garden productivity or colony growth), as predicted by symbiont-choice theory. Received 24 February 2006; revised 23 June 2006; accepted 26 June 2006.     

Influence of Experience on the Response of Bathyplectes curculionis (Hymenoptera: Ichneumonidae), a Nonaphidophagous Parasitoid, to Aphid Odor

Bathyplectes curculionis (Thomson) is an introduced natural enemy of the alfalfa weevil in North America. The wasp requires carbohydrate foods as an adult. Adult wasps have increased longevity and fecundity when provided access to pea aphid, Acyrthosiphon pisum (Harris), honeydew in the laboratory, and adults respond positively to the presence of pea aphids in alfalfa fields. However, it is unknown how these wasps find aphid honeydew in the field. In a series of Y-tube olfactometer experiments, we evaluated the response of naïve and experienced adult female B. curculionis to odors from pea aphids, alfalfa, and pea aphids on alfalfa. Naïve adult females did not respond positively to pea aphid odor even when hungry. But adult females were able to learn aphid odor, and the mechanism of learning appears to be associative rather than by sensitization. Naïve females also showed no preference for alfalfa odor but learned alfalfa odor through sensitization. The wasps did not distinguish between alfalfa with aphids and alfalfa without aphids, even after exposure to aphids or alfalfa with aphids. However, they preferred pea aphid odor to alfalfa odor after a feeding experience in the presence of pea aphid odors. But after exposure to mixed odors of aphids and alfalfa while feeding, B. curculionis females preferred the odor of alfalfa to the odor of pea aphids. These results suggest that alfalfa odors mask or override aphid odors when aphids are associated with alfalfa (as happens naturally), thus interfering with the wasp's ability to respond to learned aphid odors. Therefore, although the wasps are capable of learning to find pea aphids and their honeydew in a simplified laboratory setting, it appears unlikely that they do so in the field.     

Nest-area marking with faeces: a chemical signature that allows colony-level recognition in seed harvesting ants (Hymenoptera, Formicidae)

Summary. Nestmate recognition systems in ants are largely based on chemical signals. The hydrocarbon fraction of the lipid layer which covers the insect cuticle plays a determinant role in this context. Here we report a novel extension of nestmate and alien recognition – nest area marking with faeces containing the same hydrocarbons as the cuticle of workers – in a harvesting ant, Messor capitatus. Workers of M. capitatus deposit large quantities of brown-yellow material from the hindgut (termed spots) in the vicinity of the nest. Behavioural investigation showed that such spotting behaviour has a communicative value in the context of nest area identification. Anal fluids deposited in the nest surroundings contain colony-specific cues which the ants use to recognize their own nest areas, and distinguish them from foreign areas even in the absence of nestmate or alien ants. Chemical analyses by gas chromatography-mass spectrometry (GC-MS) of the contents of anal spots, rectal sacs, and cuticular extracts revealed that all contain the same long-chained linear and branched hydrocarbons in varying proportions. Importantly, multivariate analyses showed that the relative proportions of these compounds on the cuticle and in spots are colony-specific. This provides a mechanism by which spot marking could be used by workers to define and recognize their colony area, and would represent a simple extension of the existing nestmate recognition template based on colonial cuticular signatures. The ecological and sociobiological implications of these findings are discussed.Received 3 February 2004; revised 10 June 2004; accepted 14 June 2004.     

Effects of dietary sucrose concentration on aphid honeydew carbohydrate levels and rates of excretion

Honeydew droplets freshly excreted by two aphid species, Myzus persicae (Sulz.) and Acyrthosiphon pisum (Harr.), feeding on nutritionally adequate, artificial diets containing 10–40% w/v sucrose, were collected in oil. The volume, frequency of excretion, and carbohydrate content of the droplets were determined by measuring the diameter of the droplets in the oil, by recording the number of droplets excreted per aphid per hour, and by using a quantitative colorimetric method for estimating carbohydrates. Rates for the intake or excretion of fluid, the intake and absorption of sucrose, and the excretion of carbohydrate were consequently calculated for the two species on each dietary sucrose concentration. The percentage of dietary sucrose absorbed by the aphids ranged from 36% on diet with 40% sucrose to 97% on diet with 10% sucrose. Such honeydew/diet relationships may allow one to use honeydew carbohydrate concentrations to estimate the sucrose concentrations of the phloem sap ingested by aphids feeding on host plants.

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Résumé Des gouttelettes de miellat fraîchement excrétées par deux espèces de pucerons, Myzus persicae (Sulz.) et Acyrthosiphon pisum (Harr.), pucerons nourris sur des régimes nutritifs synthétiques contenant 10–40% de saccharose, ont été récupérées dans de l'huile. Le volume des gouttelettes a été déterminé en mesurant leur diamètre individuel dans l'huile, la fréquence d'excrétion établie par le nombre de gouttelettes excrétées par puceron par heure, et la teneur en glucides par une méthode d'analyse colorimétrique quantitative. Les taux d'ingestion et d'excrétion de liquide, d'ingestion et d'absorption de saccharose ainsi que les taux d'excrétion de glucides ont été par la suite calculés pour les deux espèces d'aphides nourris sur les différentes concentrations de saccharose. Le pourcentage de saccharose alimentaire absorbé par les aphides a été de 36% sur le régime contenant 40% de ce glucide, mais s'est élevé à 97% sur le régime ne contenant que 10% de saccharose. D'après ces relations miellat/régime nutritif, et en connaissant la teneur du miellat en glucides, on peut faire une approximation de la concentration du saccharose dans la sève élaborée du phloème ingérée par les aphides.

     

Disruption of parasitism of the California red scale (Homoptera: Diaspididae) by three ant species (Hymenoptera: Formicidae)

The relative influence of the southern fire ant, Solenopsis xyloni McCook, Argentine ant, Linepithema humile (Mayr), and native gray ant, Formica aerata (Francoeur), on parasitism of California red scale, Aonidiella aurantii Maskell, was studied in the laboratory for two parasitoids, Comperiella bifasciata Howard and Aphytis melinus DeBach. All three ant species reduced percentage parasitism by C. bifasciata and both percentage parasitism and host mutilation by A. melinus. Southern fire ant was the least disruptive and native gray ant the most disruptive. Southern fire ant removed 12% of scale from the lemons, presumably to feed on them, while the other ant species did not exhibit significant removal of scale compared to the controls. Percentage parasitism of California red scale exhibited by C. bifasciata was more than 2-fold the level exhibited by A. melinus. Percentage mutilation of California red scale, including probing and host feeding, was nearly 5-fold higher for A. melinus than C. bifasciata. Because A. melinus required a longer total host examination + oviposition period in the absence of ants than C. bifasciata and because oviposition occurs as the last act in a sequence of behaviors, disruption by the ants had a more significant negative effect on oviposition by A. melinus.     

Pathogenicity of fungal isolates (Ascomycota: Hypocreales) against Peregrinus maidis, Delphacodes kuscheli (Hemiptera: Delphacidae), and Dalbulus maidis (Hemiptera: Cicadellidae), vectors of corn diseases

Preliminary screening assays were carried out on 17 isolates from five fungal species Beauveria bassiana, Lecanicillium muscarium, Metarhizium anisopliae, Isaria farinosa, and I. fumosorosea. The three most effective isolates against Peregrinus maidis (Hemiptera: Delphacidae) were B. bassiana CEP 147, CEP 150, and CEP 189. There were no consistent differences found in males and females regarding fungal susceptibility. However, more females than males were proportionally infected. There was not a correlation between the percentage of conidial germination and the percentage of mortality caused by fungal infection in any of the treatments. Only B. bassiana CEP 147, which caused a cumulative mortality of 69.8 ± 6.4% after 7 days post-inoculation, was selected to be assayed against adults of P. maidis, Delphacodes kuscheli (Hemiptera: Delphacidae), and Dalbulus maidis (Hemiptera: Cicadellidae). In pathogenicity tests significant differences were observed among treatments. After 2 weeks post-inoculation, both D. kuscheli (cumulative mortality of 73.3 ± 9.0%) and P. maidis (cumulative mortality of 68.6 ± 6.7%) were significantly more susceptible than D. maidis (cumulative mortality of 49.9 ± 9.7%) to the selected isolate.     

Distribution and Abundance of Mymarid Parasitoids (Hymenoptera: Mymaridae) of Sophonia rufofascia Kuoh and Kuoh (Homoptera: Cicadellidae) in Hawaii

The abundance of mymarid parasitoids attacking the two-spotted leafhopper (Sophonia rufofascia [Kuoh and Kuoh]), a polyphagous pest recently adventive to Hawaii, was monitored using yellow sticky cards deployed in several areas on the islands of Kauai and Hawaii. The yellow cards captured Chaetomymar sp. nr bagicha Narayanan, Subba Rao, & Kaur and Schizophragma bicolor (Dozier), both adventive species, and Polynema sp. Haliday, which is endemic to Hawaii (Hymenoptera: Mymaridae). The former two species were most abundant at all sites. On Kauai, there was a negative correlation between the captures of C. sp. nr bagicha and those of Polynema sp. Throughout the season, the increase in parasitoid numbers generally followed the increase in leafhopper numbers. C. sp. nr. bagicha and S. bicolor showed distinct habitat preferences. Removal of Myrica faya Aiton, an invasive weed that is a highly preferred two-spotted leafhopper host, decreased the overall numbers of captured parasitoids, but led to a twofold increase in the ratio of trapped parasitoids/hosts in weed-free areas.     

Lineage specific evolution of an alternative social strategy in Tetramorium ants (Hymenoptera: Formicidae)

The western Palearctic ant, Tetramorium moravicum , is the only species of the large genus known to display two alternative social strategies: macrogyny-monoygyny (characterized by one large queen per colony) and microgyny-polygyny (multiple small queens per colony). The microgyne colonies of this ant are confined to the westernmost part of its distribution area and occasionally co-occur with macrogyne colonies. We investigated phylogenetic and geographical patterns in relation to social strategy by analysing 1031 bp of the mitochondrial cytochrome c oxidase subunit I gene of T. moravicum from the entire known species range. Phylogenetic analyses revealed two clear-cut lineages coinciding with the eastern and western populations, which probably were separated after the late Quaternary glacial periods. Geographical distance analysis suggested allopatric fragmentation of the two lineages. We hypothesize that the two lineages colonized Central Europe postglacially from two directions. The current distribution gap of 500 km between the tip of the eastern lineage, in the Czech Republic, Austria and Hungary, and the western lineage, in Germany, was confirmed by hierarchical population analysis. Macrogyny-monogyny probably was the original condition in T. moravicum ; the alternative strategy of microgyny-polygyny may have evolved at the tip of the western lineage, as recently as the early Holocene.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 247–255.     

Labial gland disease in the genusFormica (Formicidae,Hymenoptera)

Summary Labial gland disease causes swelling of the labial glands in the pupal stage inFormica species. The resulting adults have enlarged thoraces and are called secretergates (Wasmann's pseudogynes). They infect the younger larvae, probably during feeding. The agent of the disease is unknown. It soon becomes non-infectious outside the labial glands. Diseased females were not seen to feed larvae. Yet their offspring contained secretergates. The discrepancy remains unexplained.     

Aphis clerodendri Matsumura (Hemiptera: Aphididae), attendant ants (Hymenoptera: Formicidae) and associates on Clerodendrum (Verbenaceae) in Australia

Abstract Aphis clerodendri Matsumura is newly recorded from Australia and is known from the Northern Territory, on islands in Torres Strait, and in rainforest in northern Queensland and New South Wales. It induces the formation of leaf pseudogalls on native species of Clerodendrum and is commonly attended by ants, which penetrate and may polydomously nest in the galls. Previously known only from eastern Asia, A. clerodendri can now be classified as native to Australia and Australasian in natural distribution. The species is also newly recorded from Papua New Guinea and Vietnam.     

Density-dependent predation of the antLasius niger (Hymenoptera: Formicidae) on two attended aphidsLachnus tropicalis andMyzocallis kuricola (Homoptera: Aphididae)

The antLasius niger was observed collecting honeydew and preying on the two aphid speciesLachnus tropicalis andMyzocallis kuricola on the chestnut treesCastanea crenata. Observation determined how the antL. niger controlled their predation on the aphids in response to the density and honeydew-productivity of the aphids.Lachnus tropicalis was a better honeydew source thanM. kuricola forL. niger in terms of the amount of honeydew collected per unit time by the ants. The number of foraging workers on a tree increased with the number ofL. tropicalis on the tree, but not with the number ofM. kuricola. The density ofL. tropicalis perL. niger worker on a tree had a positive effect on the predation activity ofL. niger on both aphids, whereas the density ofM. kuricola per ant did not have any significant effect. The predation pressure by the ant which increased withL. tropicalis density, however, directed toM. kuricola rather than toL. tropicalis. These facts suggest (1) thatL. niger control their predation activities on aphids with regards to the densities of the attended aphids per worker, and (2) that the ants prey on the aphid species producing less honeydew. The effects of the ant predation on aphids and the importance of these predation effect in antaphids interactions were discussed.