Behavior of red wood ants (Hymenoptera,Formicidae) during interaction with different symbiont partners

A comparative analysis of the behavior of Formica polyctena Först during interaction with different symbionts (free-living aphids Aphis grossulariae Kalt. and hidden larvae of the sawfly Blasticotoma filiceti Klug) was carried out. Red wood ants demonstrate different levels of functional differentiation in relatively constant groups of foragers collecting honeydew. A deep “professional” specialization with clear division of a number of tasks among foragers was studied in groups of ants tending aphids. Four professional groups of foragers with different tasks were revealed: “shepherds,” “guards,” “transporters,” and “scouts” (or “coordinators”). The groups of foragers caring for sawfly larvae mainly consist of unspecialized ants. Only few ants (about 5%) remain on duty on the fern plant near B. filiceti larvae and protect the food resource from competitors, especially from other ants. In addition, the ants demonstrate simpler behavior while collecting the larval excretion, resembling that at the sugar feeders. On the whole, the behavior of red wood ants is rather flexible. The level of functional differentiation in groups of foragers collecting honeydew is determined not only by the colony size and requirements but by the nature of their interaction with trophobionts, particularly, by the possibility of direct contact.     

Consequences of the arms race between <Emphasis Type="Italic">Maculinea teleius</Emphasis> social parasite and <Emphasis Type="Italic">Myrmica</Emphasis> host ants for myrmecophilous butterfly conservation

The arms race between Maculinea butterflies and Myrmica host ants leads to local host-parasite adaptations. In our study, we assessed whether sympatric and allopatric Myrmica scabrinodis populations exhibit behavioural differences towards Maculinea teleius larvae during the adoption-period when butterfly larvae need to be taken inside the Myrmica nest. The second aim was to assess the butterfly survival rate inside ant colonies from different populations. We used one sympatric host population and three allopatric populations: one infested by M. teleius and two uninfested populations. We found that ants from the sympatric population showed a higher number of positive behaviours toward M. teleius larvae during adoption than ants from the allopatric populations. There were no differences in the number of inspection or negative behaviour events. The survival of butterfly larvae was highest inside sympatric host colonies and differed from the survival of M. teleius reared by ants from the allopatric, uninfested populations. No difference was found for the survival rate of M. teleius raised by infested, allopatric host colonies compared to sympatric host populations. Our results suggest the lack of behavioural counter-adaptations of local hosts of M. teleius that more easily adopt and rear butterfly caterpillars compared to naive M. scabrinodis colonies. Our results may also have implications for Maculinea butterfly conservation, especially for reintroduction programmes. We suggest that the existence of behavioural host defences should be checked for the source host population, as well as for the Myrmica population from the reintroduction site. It may also be reasonable to introduce several Myrmica host colonies from the source butterfly host population.     

The importance of resource distribution: spatial co-occurrence of host plants and host ants coincides with increased egg densities of the Dusky Large Blue <Emphasis Type="Italic">Maculinea nausithous</Emphasis> (Lepidoptera: Lycaenidae)

The occurrence of the Dusky Large Blue Butterfly (Maculinea nausithous) critically depends on the availability of two key resources: the Great Burnet (Sanguisorba officinalis) as primary nectar source for adults, for egg laying and early larval development, and the host ant Myrmica rubra as the food of late instar larvae. Thus, their distributions are key parameters shaping habitat suitability, and we expected that overlapping of both resources would have a strong impact on the size of local M. nausithous populations. Their egg density may be affected (a) by the fraction of host plants per site located within My. rubra activity ranges at the patch scale, or (b) by the availability of host plants with host ants in close range at the local scale, due to the potential ability of butterfly females to detect their host ants. To test the above hypothesis, we recorded spatial distribution patterns of host plants and host ants on 29 study sites in south-western Germany and related them to egg density data of M. nausithous. We found a positive relationship between co-occurence of host plant and host ant and M. nausithous egg density at the patch scale, whereas no correlation was found at the local scale. Thus, focal populations are strongly limited by the abundance of host plants, covered with My. rubra activity, as ant-mediated oviposition could not be proved. Our results underline the importance of resource distribution; the understanding of its impacts may provide useful insights into how M. nausithous habitats can be managed in order increase their carrying capacity.     

Behavioral strategy of a lycaenid (Lepidoptera) caterpillar against aggressive ants in a Brazilian savanna

Myrmecophily is widespread in lycaenid butterflies, in which ants receive food resources and, in turn, protect caterpillars against natural enemies. This interaction ranges from obligate myrmecophily, in which immatures are invariably associated with ants and are dependent on ants for survival, and facultative myrmecophily, in which larvae are not dependent on ants for survival, but the presence of the latter may increase larvae survival. Lycaenids also include non-myrmecophilous butterflies, which do not have positive associations with ants and have developed strategies to avoid being attacked or preyed upon by them. In this study, we examined the relationship between the lycaenid Michaelus ira and two ant species associated with Distictella elongata (Bignoniaceae). This plant has extrafloral nectaries and is patrolled by Camponotus crassus and Ectatomma tuberculatum. Morphological analyses revealed that M. ira larvae have ant organs, such as dorsal nectary organs and perforated cupolas, structures associated with myrmecophily. We performed larval exposure experiments in the field, predicting that, in the absence of myrmecophily, the butterfly larva would present strategies to avoid ant attack. Results showed that larvae were attacked by both ant species. To escape ant molestation, larvae lived and fed inside silk-sealed D. elongata flower buds. We concluded that the M. ira bud-sheltering behavior was a defensive strategy against these ant species, while the dorsal nectary organs were apparently nonfunctional. Nonetheless, myrmecophily, in general, cannot be excluded in M. ira since relationships with other ant species may exist.     

Parasitoids (Hymenoptera,Eulophidae) of sawflies of the genera <Emphasis Type="Italic">Fenusa,Pontania</Emphasis>, and <Emphasis Type="Italic">Arge</Emphasis> (Hymenoptera,Symphyta) from the middle Volga area

In the middle Volga area, four sawfly species (Fenusa ulmi, F. pusilla, Pontania proxima, and Arge ochropus) are attacked by 10 parasitoid species of the family Eulophidae. Of these eulophid species, seven are recorded as sawfly parasitoids for the first time. Arge ochropus is for the first time recorded as a host of Elahertus gallicus.     

Dynamics of host plant selection and host-switching by silver-spotted skipper caterpillars

Investigations of host plant selection in herbivorous arthropods have emphasized the importance of oviposition site selection by adults; however, a more complete picture of this process requires additional consideration of the factors influencing host plant choice during the immature feeding stages. We conducted a series of larval choice experiments to examine both the innate and induced preferences of larvae of the Silver-spotted skipper (Epargyreus clarus L.) on three commonly used hosts (Wisteria, Robinia, and Pueraria). Late instar E. clarus larvae reared on each of the three host plants displayed an overall pattern of innate preferences that correlated well with larval performance measures and reflected differences in foliar nutrient concentrations. Larval preferences were also influenced by rearing host species, indicating a role for feeding-induced preferences. When larvae reared on low-quality Wisteria for the first four instars were switched to higher quality Pueraria for the final instar, they developed more quickly and attained significantly higher pupal mass than larvae maintained on Wisteria throughout development. Similarly, larvae switched from Pueraria to Wisteria for the final instar suffered increased development time and produced significantly smaller pupae than those maintained on Pueraria throughout. Thus host-switching, particularly during the more mobile final instars, appears to offer larvae an opportunity to recoup fitness losses associated with early development on a low-quality host. For an equal amount of consumption, larvae feeding on Pueraria gained 50% more mass than those feeding on Wisteria, reflecting measured differences in foliar nitrogen concentration; despite these overall differences in quality, larval growth efficiency was similar among hosts. Especially in the age of common exotic plant introductions, a full understanding of the behavioral component of host selection by herbivorous insects requires appreciation of the dynamic role that immatures can play in host selection and use.     

Host plants influence female oviposition and larval performance in West Indian sweet potato weevils <Emphasis Type="Italic">Euscepes postfasciatus</Emphasis> (Coleoptera: Curculionidae)

Euscepes postfasciatus (Fairmaire) is an invasive pest of the sweet potato (Ipomoea batatas) and is also parasitic to other wild host plants of the Ipomoea genus. The population density of E. postfasciatus is sometimes greater in Ipomoea pes-caprae L. than in Ipomoea indica (Burm. f.). We investigated the desirability of I. pes-caprae as a host plant for E. postfasciatus in terms of reproductive and developmental potential. Females laid fewer eggs on I. pes-caprae, and the eclosion of their larvae was delayed compared with on I. indica. Furthermore, the larval growth rate was slower on I. pes-caprae than on I. indica. These results suggest that I. pes-caprae is not always the preferred host for egg laying and growth rate in the early developmental stages. However, the larval survival rate after the initial period of development was markedly better on I. pes-caprae than on I. indica. The present simulation study demonstrated that the population density of E. postfasciatus on I. pes-caprae overwhelmed that on I. indica over generations. Comparing the two wild host plant species, I. pes-caprae outweighs I. indica with respect to total population growth, but reproduction on I. indica may be advantageous for the colonization of the new habitat.     

Attraction of the Larval Parasitoid <Emphasis Type="Italic">Spintherus dubius</Emphasis> (Hymenoptera: Pteromalidae) to Feces Volatiles from the Adult <Emphasis Type="Italic">Apion</Emphasis> Weevil Host

The behavioral response of the larval parasitoid Spintherus dubius (Hymenoptera: Pteromalidae) to volatile compounds derived from its Apion weevil hosts was investigated in two-choice bioassays. Odor source candidates were the larval and adult stages of weevils, clover flowers, and feces from adult weevils. Despite S. dubius being a larval parasitoid, the odor of weevil larvae isolated from the clover flowers was not attractive to female parasitoids. Surprisingly, S. dubius females were instead attracted by the odor from the feces of adult weevils. The female parasitoids were similarly attracted to the feces produced by the two main hosts, the red clover weevil (A. trifolii) and the white clover weevil (A. fulvipes). Chemical analysis of the volatile composition of feces produced by the two hosts revealed qualitatively similar odor profiles, correlating with the observed attraction by the parasitoid towards both odor sources. Some of the identified volatile compounds are commonly present in clover plant headspace fractions and may function as a kairomone to facilitate orientation by S. dubius to Apion-infested clover flowers. Larval and adult weevils were not attractive for parasitoid females, whereas, for the white clover weevil-plant association, infested flowers were highly attractive. These data show the use by the clover weevil parasitoid of an alternative source of olfactory information for locating its host.     

Larvae of the exotic predatory ladybird <Emphasis Type="Italic">Platynaspidius maculosus</Emphasis> (Coleoptera: Coccinellidae) on citrus trees: prey aphid species and behavioral interactions with aphid-attending ants in Japan

An aphidophagous ladybird, Platynaspidius maculosus (Weise) (Coleoptera: Coccinellidae), is originally distributed in China, Taiwan, and Vietnam. The ladybird has recently intruded into the southern and central parts of Japan. The present study found that the larvae of this ladybird preyed on three aphid species, Aphis spiraecola, Aphis gossypii, and Toxoptera citricidus (all Hemiptera: Aphididae), feeding on young shoots of various Citrus species in August to early October in Shizuoka Prefecture, central Japan. Laboratory rearing of the sampled larvae confirmed that the larvae completed their development (adult emergence) by consuming each of the three aphid species. The ladybird larvae were observed foraging in aphid colonies attended by one of the four ants, Lasius japonicus, Pristomyrmex punctatus, Formica japonica, and Camponotus japonicus (all Hymenoptera: Formicidae). Field observations revealed that the foraging/feeding larvae were almost completely ignored by honeydew-collecting ants even when they physically contacted each other. Thus, in Japan, the larvae of the exotic ladybird exploit colonies of the three aphid species attended by one of the four ant species on many Citrus species. On the basis of the results, I discuss the possibility of the ladybird’s reproduction on citrus trees in Japan, probable adaptations of the ladybird larvae to aphid-attending ants, and potential impacts of the ladybird on native insect enemies attacking ant-attended aphids on citrus.     

Ant-partners play a minor role on occurrence of the myrmecophilous butterfly <Emphasis Type="Italic">Leptotes cassius</Emphasis> in its host plant

Ant-related oviposition in facultatively myrmecophilous lycaenid butterflies is common, but not universal. In fact, our knowledge of ant-related oviposition in lycaenids is based on some common species (e.g., Rekoa marius, Allosmaitia strophius, Parrhasius polibetes), which limits generalizations about these systems. In this study, we experimentally investigated whether the oviposition pattern of the florivorous lycaenid Leptotes cassius was influenced by the presence of Camponotus ants and whether larvae were attended, rather than attacked, by ants. This might be evidence of myrmecophily. Both L. cassius and Camponotus ants occur on Bionia coriacea, an extrafloral nectaried legume shrub that grows in the Brazilian cerrado. Plants were randomly assigned to ant-present and ant-excluded treatments and were observed twice throughout the short reproductive season. Larvae of L. cassius were tended by ants, whose attendance was characterized by active antennation on the last body segments of the caterpillars. Therefore, Camponotus can be considered a partner of L. cassius. Lycaenid abundance was on average 1.9- and 1.21-fold higher in plants with ants in each sampling period, respectively, indicating a tendency of L. cassius to occur in plants with ants. Nonetheless, results were not statistically significant, suggesting that in this case ants are not a major cue for lycaenid oviposition. In many ant–lycaenid mutualisms, butterfly immatures benefit from reduced parasitism rates. However, no L. cassius immature, regardless of ant presence or absence, was parasitized. Furthermore, larvae may occur inside flower buds that may provide protection from natural enemies; thus, ants may not be required for immature protection.     

The Predation Strategy of the Recluse Spider <Emphasis Type="Italic">Loxosceles rufipes</Emphasis> (Lucas, 1834) against four Prey Species

Generalist predators have to deal with prey with sometimes very different morphologies and defensive behaviors. Therefore, such predators are expected to express plasticity in their predation strategy. Here we investigated the predatory behavior of the recluse spider Loxosceles rufipes (Araneae, Sicariidae) when attacking prey with different morphologies and defensive mechanisms. We expected L. rufipes to show different prey capture strategies and variable acceptance towards each prey type. Potential prey species were collected directly from the web or in the surroundings of the web-building site of L. rufipes. We collected and used the following in our experiments: termite workers (Nasutitermes sp.), lepidopteran larvae (Eurema salome), ants (Camponotus sp.) and isopods (Tylidae). We paired these prey with L. rufipes and recorded their behavior in captivity, quantifying acceptance rate, immobilization time and the sequence of behaviors by the predator. The acceptance rate was lower for isopods but not different among other prey. The immobilization time was higher for isopods than for termites and similar for the other pairwise comparisons. The behavioral sequence was similar for all prey except for isopods, which were also bit more often. Our combined results show plasticity in the behavior of L. rufipes and also show it subdues a potentially dangerous prey (ant) and an armored prey (isopod).     

On the Adult Behavioral Repertoire of the Sawfly <Emphasis Type="Italic">Perreyia flavipes</Emphasis> Konow, 1899 (Hymenoptera: Pergidae): Movement,Mating, and Thanatosis

Perreyia flavipes Konow, 1899 (Pergidae: Perreyinae) is a sawfly species with gregarious larvae commonly found in open areas in southern Brazil through Uruguay and northern Argentina. The combination of highly gregarious larvae with the production of a variety of toxins in P. flavipes has led to severe cases of intoxication in a variety of livestock species. Over the years, considerable information was discovered on the larval natural history of P. flavipes; however, virtually nothing is known about the adult behavior, particularly because of its short life-span. In this study, we report on the first extensive study on the adult behavior of P. flavipes, including movement, mating (with oviposition and maternal care), and thanatosis. Our results show some unusual behavioral adaptations presented by P. flavipes, such as irregular activity behavior (i.e., deficient gait pattern), thanatosis behavior-like display, and primitive maternal care. Individual behavioral acts are described and compared among sexes, and their potential functions are discussed.     

On distribution and ecology of some species of cleonines (Coleoptera,Curculionidae): IV. Genus <Emphasis Type="Italic">Lixus</Emphasis> F., subgenus <Emphasis Type="Italic">Eulixus</Emphasis> Reitt.

Data on habitats, food plants of larvae and adults, feeding, mating, oviposition, larval and pupal development, natural enemies, and distribution of seven weevil species (Lixus canescens F.-W., L. iridis Ol., L. myagri Ol., L. punctirostris Boh., L. subtilis Boh., L. incanescens Boh., and L. brevipes Bris.) are given. New host plants of L. canescens and L. iridis are revealed. Gall induction by L. brevipes is reported for the first time. Distribution of all the species in Ukraine and Russia (the latter based on the literature) are given in more detail. Information on the known and potential economic importance of every species is provided.     

Host ant use of the Alcon blue butterfly at the northern range margin

In myrmecophilous insects, interactions with ants are often a key factor determining persistence of their populations. Regional variation in host ant use is therefore an essential aspect to consider to provide adequate conservation practices for such species. In this study, we examined this important facet of species’ ecology in an endangered myrmecophilous butterfly Phengaris (=Maculinea) alcon (Lepidoptera, Lycaenidae). The investigations conducted in peripheral populations in Estonia allowed us to expand the knowledge of its host ant use to the northern distribution limit of the species. Our data indicate that in its northernmost populations, the xerophilous ecotype of Phengaris alcon is primarily parasitizing a single host ant species, Myrmica schencki. The data collected are in line with the emerging evidence suggesting that peripheral and core populations of P. alcon use different host ants, and peripheral populations tend to display higher host ant specificity. We also show that, at its northern range margin, P. alcon might be more limited by the availability of its sole larval food plant in the region, Gentiana cruciata, than the densities of its host ant. Finally, we found a strong negative correlation between Myrmica spp. and Lasius spp. colony densities, suggesting that interspecific competition between ants could have a substantial influence on host ant availability of Phengaris butterflies, and thus should be taken into account in conservation plans of these species.     

Host plant quantitative trait loci affect specific behavioral sequences in oviposition by a stem-mining insect

Key message

Genetic diversity in quantitative loci associated with plant traits used by insects as cues for host selection can influence oviposition behavior and maternal choice.

Abstract

Host plant selection for oviposition is an important determinant of progeny performance and survival for phytophagous insects. Specific cues from the plant influence insect oviposition behavior; but, to date, no set of host plant quantitative trait loci (QTLs) have been shown to have an effect on behavioral sequences leading to oviposition. Three QTLs in wheat (Triticum aestivum L.) have been identified as influencing resistance to the wheat stem sawfly (WSS) (Cephus cinctus Norton). Wheat near-isogenic lines (NILs) for each of the three QTLs were used to test whether foraging WSS were able to discriminate variation in plant cues resulting from allelic changes. A QTL on chromosome 3B (Qss-msub-3BL) previously associated with stem solidness and larval antibiosis was shown to affect WSS oviposition behavior, host preference, and field infestation. Decreased preference for oviposition was also related to a QTL allele on chromosome 2D (Qwss.msub-2D). A QTL on chromosome 4A (Qwss.msub-4A.1) affected host plant attractiveness to foraging females, but did not change oviposition preference after females landed on the stem. These findings show that oviposition decisions regarding potential plant hosts require WSS females to discriminate signals from the plant associated with allelic variation at host plant quantitative loci. Allele types in a host plant QTL associated with differential survival of immature progeny can affect maternal choices for oviposition. The multidisciplinary approach used here may lead to the identification of plant genes with important community consequences, and may complement the use of antibiosis due to solid stems to control the wheat stem sawfly in agroecosystems.

     

Defensive Responses in Groundnut Against Chewing and Sap-Sucking Insects

Induced resistance is one of the important components of host plant resistance to insects. We studied the induced defensive responses in groundnut genotypes with different levels of resistance to the leaf defoliator Helicoverpa armigera and the sap-sucking insect Aphis craccivora to gain an understanding of the induced resistance to insects and its implications for pest management. The activity of the defensive enzymes (peroxidase, polyphenol oxidase, phenylalanine ammonia lyase, superoxide dismutase, ascorbate peroxidase, and catalase) and the amounts of total phenols, hydrogen peroxide, malondialdehyde, and proteins were recorded at 6 days after infestation. Induction of enzyme activities and the amounts of secondary metabolites were greater in the insect-resistant genotypes ICGV 86699, ICGV 86031, ICG 2271, and ICG 1697 infested with H. armigera and A. craccivora than in the susceptible check JL 24. The resistant genotypes suffered lower insect damage and resulted in lower Helicoverpa larval survival and weights than those larvae fed on the susceptible check JL 24. The number of aphids was significantly lower on insect-resistant genotypes than on the susceptible check JL 24. The results suggested that groundnut plants respond to infestation by H. armigera and A. craccivora in a similar way; however, the degree of the response differed across the genotypes and insects, and this defense response is attributed to various defensive enzymes and secondary metabolites.     

Host-plant patch qualities and presence of a likely competitor species affect the distribution and abundance of a rare British moth, <Emphasis Type="Italic">Cucullia lychnitis</Emphasis>

Understanding the environmental factors that affect the distribution and abundance of rare and threatened species can help identify priority sites for conservation action such as habitat management. The moth Cucullia lychnitis is identified in UK legislation as a priority species for conservation. Its conspicuous larvae feed on a patchily distributed host plant Verbascum nigrum, but are entirely absent from some host plant patches within the species’ range. Environmental variability among patches was investigated to understand factors affecting (i) patch occupancy by C. lychnitis and (ii) abundance of C. lychnitis, where present. Occupancy of individual V. nigrum plants within occupied patches was also investigated. The likelihood of patch occupancy increased with patch size, and decreased with patch isolation. Abundance of C. lychnitis was negatively correlated with patch isolation and with abundance of the weevil Cionus nigritarsis, which exploits the same host plant and may be a competitor. Within occupied patches, larvae of C. lychnitis were significantly more likely to be found on taller plants with more flower spikes; such plants are typical of established rather than newly-created patches. These results may help to guide efforts to conserve C. lychnitis, highlighting the importance of habitat connectivity and indicating potential management actions to promote features positively associated with larval incidence and abundance.     

Pathogen invasion triggers an evolutionary trap for an endangered checkerspot butterfly dependent on an exotic host plant

Exotic pathogen invasions can change host eco-evolutionary interactions and possibly create an evolutionary trap when the pathogen generates mismatches between developmental phenology and reproductive cues. Taylor’s checkerspot butterfly (Euphydryas editha taylori), is an endangered species of western North America with 80 % of the extant populations dependent on an exotic host, Plantago lanceolata. Survey of occupied, recently extinct, and unsuccessful butterfly reintroduction sites spanning 4° of latitude revealed widespread disease on P. lanceolata caused by Pyrenopeziza plantaginis. This fungal pathogen, new to North America, reduces the standing crop of P. lanceolata foliage throughout the winter post-diapause larval feeding period. However, disease is absent when adult butterflies and pre-diapause larvae are active. Diseased plants were frequent in Taylor’s checkerspot populations with a history of persistence, but >90 % of the host plants in these populations had initiated new leaves within the first few weeks of post-diapause larval feeding. Conversely, host plants in recently extinct and unsuccessfully reintroduced populations were severely diseased, >66 % mean foliage necrosis/plant, and <23 % had initiated new leaves. Feeding choice trials with 25 larvae indicated that new leaves were strongly and consistently preferred by post-diapause larvae over all other available leaf types, both diseased and non-diseased. Because the influence of disease on post-diapause larval food resources is developmentally disassociated from oviposition, P. plantaginis invasion appears to have triggered an evolutionary trap for Plantago-dependent populations of Taylor’s checkerspot.