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      The Exocrine Chemistry of the Parasitic Wasp Sphecophaga orientalis and Its Host Vespa orientalis: A Case of Chemical Deception?

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          Abstract

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          The wasp Sphecophaga orientalis is a parasitoid of the Oriental hornet ( Vespa orientalis) in its subterranean colonies. We describe this parasitoid’s occurrence in hornet nests in Israel and compare the chemical composition of cuticular washes of both species. The dissimilarity between the two excludes the possibility that the parasite uses either camouflage or chemical mimicry to evade host aggression. Because the parasitoid features large amounts of the necrophoric compound oleic acid, we suggest that, due to this compound, the host considers the parasite as refuse and ignores its presence. The parasitoid head also contains rose oxide, a repellent, possibly used to repel aggressive workers and which, combined with its necrophoric odor, enables it to remain in the nest undisturbed.

          Abstract

          The main challenge facing a parasite of social insects lies in deceiving its host’s detection and defense systems in order to enter and survive within the host colony. Sphecophaga orientalis is an ichneumonid wasp that parasitizes the pupae of the Oriental hornet Vespa orientalis. In Israel’s Mediterranean region, this parasitoid infects on average 23.48% (8–56%) of the host pupal cells. Observation of colonies brought to the laboratory revealed that the parasite moves around within the colony without being aggressed by the host workers. To assess how the parasite evades host detection and defense, we compared the cuticular hydrocarbon (CHC) profiles of both species. There was little similarity between the parasite and the host workers’ CHC, refuting the hypothesis of chemical mimicry. The parasite’s CHCs were dominated by linear alkanes and alkenes with negligible amounts of branched alkanes, while the host workers’ CHCs were rich in branched alkanes and with little or no alkenes. Moreover, the parasite cuticular wash was markedly rich in oleic acid, previously reported as a cue eliciting necrophoric behavior. Since nests of Oriental hornets are typified by large amounts of prey residues, we suggest that, due to its unfamiliar CHCs and the abundance of oleic acid, the parasite is considered as refuse by the host. We also detected rose oxide in the parasitoid head extracts. Rose oxide is a known insect repellent, and can be used to repel and mitigate aggression in workers. These two factors, in concert, are believed to aid the parasite to evade host aggression.

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          Chemical ecology and social parasitism in ants.

          The chemical strategies by which parasites manage to break into the social fortresses of ants offer a fascinating theme in chemical ecology. Semiochemicals used for interindividual nestmate recognition are also involved in the mechanisms of tolerance and association between the species, and social parasites exploit these mechanisms. The obligate parasites are odorless ("chemical insignificance") at the time of usurpation, like all other callow ants, and this "invisibility" enables their entry into the host colony. By chemical mimicry (sensu lato), they later integrate the gestalt odor of this colony ("chemical integration"). We hypothesize that host and parasite are likely to be related chemically, thereby facilitating the necessary mimicry to permit bypassing the colony odor barrier. We also review the plethora of chemical weapons used by social parasites (propaganda, appeasement, and/or repellent substances), particularly during the usurpation period, when the young mated parasite queen synthesizes these chemicals before usurpation and ceases such biosynthesis afterwards. We discuss evolutionary trends that may have led to social parasitism, focusing on the question of whether slave-making ants and their host species are expected to engage in a coevolutionary arms race.
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            Social Immunity: Emergence and Evolution of Colony-Level Disease Protection.

            Social insect colonies have evolved many collectively performed adaptations that reduce the impact of infectious disease and that are expected to maximize their fitness. This colony-level protection is termed social immunity, and it enhances the health and survival of the colony. In this review, we address how social immunity emerges from its mechanistic components to produce colony-level disease avoidance, resistance, and tolerance. To understand the evolutionary causes and consequences of social immunity, we highlight the need for studies that evaluate the effects of social immunity on colony fitness. We discuss the role that host life history and ecology have on predicted eco-evolutionary dynamics, which differ among the social insect lineages. Throughout the review, we highlight current gaps in our knowledge and promising avenues for future research, which we hope will bring us closer to an integrated understanding of socio-eco-evo-immunology. Expected final online publication date for the Annual Review of Entomology Volume 63 is January 7, 2018. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
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              Nestmate recognition cues in the honey bee: differential importance of cuticular alkanes and alkenes.

              In social insects, recognition of nestmates from aliens is based on olfactory cues, and many studies have demonstrated that such cues are contained within the lipid layer covering the insect cuticle. These lipids are usually a complex mixture of tens of compounds in which aliphatic hydrocarbons are generally the major components. The experiments described here tested whether artificial changes in the cuticular profile through supplementation of naturally occurring alkanes and alkenes in honeybees affect the behaviour of nestmate guards. Compounds were applied to live foragers in microgram quantities and the bees returned to their hive entrance where the behaviour of the guard bees was observed. In this fashion we compared the effect of single alkenes with that of single alkanes; the effect of mixtures of alkenes versus that of mixtures of alkanes and the whole alkane fraction separated from the cuticular lipids versus the alkene fraction. With only one exception (the comparison between n-C(19) and (Z)9-C(19)), in all the experiments bees treated with alkenes were attacked more intensively than bees treated with alkanes. This leads us to conclude that modification of the natural chemical profile with the two different classes of compounds has a different effect on acceptance and suggests that this may correspond to a differential importance in the recognition signature.
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                Author and article information

                Journal
                Insects
                Insects
                insects
                Insects
                MDPI
                2075-4450
                23 December 2020
                January 2021
                : 12
                : 1
                : 2
                Affiliations
                [1 ]School of Zoology, Faculty of Life Sciences, Tel Aviv University, 6997801 Tel Aviv, Israel; dubiner@ 123456mail.tau.ac.il (S.D.); cknitzan@ 123456gmail.com (N.C.); mikavolov@ 123456mail.tau.ac.il (M.V.); hefetz@ 123456tauex.tau.ac.il (A.H.); ryaseltz@ 123456mail.tau.ac.il (R.S.)
                [2 ]Faculty of Marine Science, Ruppin Academic Center, 4025000 Michmoret, Israel
                Author notes
                [* ]Correspondence: levineran1@ 123456gmail.com
                Article
                insects-12-00002
                10.3390/insects12010002
                7822126
                33374521
                2eb9182a-e6e0-4ca1-ab44-0763b5a8e276
                © 2020 by the authors.

                Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license ( http://creativecommons.org/licenses/by/4.0/).

                History
                : 04 November 2020
                : 19 December 2020
                Categories
                Article

                vespa,sphecophaga,parasitoid,chc,oleic acid,rose oxide
                vespa, sphecophaga, parasitoid, chc, oleic acid, rose oxide

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