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Data available from the Dryad Digital Repository: In this context, we evaluated the evidence of evolutionary change in invasive populations of Harmonia axyridis Pallas Coccinellidae: We measured egg defenses—cocktail of hydrocarbons on the egg’s surface flagging egg toxicity and the concentration of the main alkaloid harmonine—in individuals from three populations along the invasion route Japan: Our results support the EICA hypothesis: The number of introduced species rocketed in the 20th Century as trade, fueled by fossil energies, went global.

In this work, we focused on the Harlequin ladybird, Harmonia axyridis Pallas Coccinellidae: Coleopteraa generalist predator of aphids Aphididae: This expansion can also derive from the recipient community’s traits anemjes environmental context.

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In the case of H. This study explores some of these processes, namely the evolution of chemical defenses. Rieder, Newbold, Sato, Yasuda, and Evans provided one of the rare comparative analyses of the H.

The authors studied the intraguild predation of the Japanese Coccinella septempunctata L. Their results indicate that the eggs from the Japanese i. These dissimilarities in egg defenses in native and invasive populations of H. However, there is a lack of comprehensive studies assessing the natural enemies of H. Enemy release can confer other advantages to the invader than just a lower mortality rate. Japan native populationUnited States of America introduced in the 80s from Asiaand South Africa presence first reported inintroduced from the United States.

Along the invasion route, we can therefore expect changes in the ECHCs profiles and in the overall quantity of ECHCs and a decrease in the concentration of harmonine in the eggs. For two generations, they were fed with ionized Ephestia kuehniella Lepidoptera: At generation G2, males and females were separated immediately after emergence to prevent mating. From then on, the adults were reared for the chemical ecology experiments as described below. Three times a week they were fed an excess of pea aphids Acyrthosiphon pisum Harris, a cosmopolitan H.

A stem of broad bean Vicia faba L. Three days later, when the cuticle hardened, the males and the females were identified. The solvent was then transferred into a 1. Whenever an egg was damaged, the whole extraction sample was discarded to avoid contamination of the egg coating with molecules from inside the eggs.

For each population, we analyzed three batches of to eggs. Following the extraction of the ECHCs, 48 to 50 eggs per population were analyzed for harmonine content.

The carrier gas was helium and the flow rate 1.


The chromatograph oven was programmed as follows: The whole system was controlled by Xcalibur data software, version 2. This nomenclature uses a descriptor XX for anmeies total number of carbons in the hydrocarbon component C XX ; i. The whole system was controlled using an Xcalibur data system, version 2.

The average mass of the eggs was calculated on samples of 48 to 50 eggs for each of the three populations. They were compared using an ANOVA after the requirements for homoscedasticity and normality had been checked for.

The total quantity of hydrocarbons on the chorion of the eggs from the anemes populations was corrected for egg mass differences before being compared using a Kruskal—Wallis test.

There were three replicates per population. This method is recommended when the number of variables exceeds the number of observations Anderson, The nMDS was based on Jaccard similarity, and the data were transformed into presence 1 or absence 0 prior to analysis. In addition, based on the ECHCs profiles, the multivariate dispersion was measured for each population using the betadisper function of the Vegan R package.

The concentration of harmonine in the eggs laid by the ladybirds of the three populations was analyzed using a GLM with a Poisson family. The three populations were compared using a priori orthogonal contrasts. All the analyses were performed using R Commander, R statistical software version 2. A total of 65 compounds were detected: The stress value is 0. There were three samples per population. On the contrary, the eggs from the more recently introduced population of South Africa contain less harmonine than the eggs from both Japan and the United States z value: Therefore, favored phenotypes can evolve allowing nonindigenous populations to spread in new environments.

Thus, we should pay more attention to the study of key phenotypic traits that might have evolved during the invasion process of H. In this study, we evaluated the evidence of evolutionary change in H.

It is generally accepted that the majority of insect hydrocarbons are de novo synthesized, and thus have fitness costs Blomquist, Therefore, we expected that enemy release should lead to a decrease in the allocation of resources to defense in favor of increased competitive ability. Our results are consistent with this hypothesis because we found a decrease in the concentration of harmonine in the eggs and a tendency toward a decrease in the number of ECHCs along the invasion route. However, we found no differences between the three populations in the total amount of ECHCs.


In insects, cuticular hydrocarbons CHCs first hemolhtiques a role in the protection against desiccation. As we found no differences in the total amount of ECHCs per egg along the invasion route, we conclude that the antidryness capacity must have been maintained in all populations. However, we found a tendency toward a decrease in the number of ECHCs as well as changes in hemoltyiques relative quantities of the different compounds along the invasion route.

The invasive populations of H. We cannot exclude the possibility that the CHC profiles of the invasive populations have evolved in the absence of selection pressure by native natural enemies.

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An unequivocal link has been shown between the harmonine content in H. Our results show that the concentration of harmonine is significantly lower in the eggs from the South African population. This could explain why we only found differences in the concentration of harmonine of the more recently introduced South African population.


Their study was carried out before anwmies, when the USA population was in an earlier phase of the invasion process. Naemies would be interesting to monitor the current South African and the native Japanese populations to see whether the differences in defense still exist. A side result, that should be noted, is the fact that in our experiments, the ladybirds of the two invasive populations laid significantly larger eggs than the native Japanese ladybirds.

Thus, we still have little understanding of how constitutive and induced chemical defenses vary among native and invasive populations of animals. Also, our observational work resulting from anemifs single, nonmanipulated situation, and based on the study of a limited number of anmeies should be taken with caution.

For a start, it would be interesting to study other H. Moreover, this study should be extended to other life stages. At last, our study is based on the premise of enemy release. Our common garden experiment thus corresponds to what Colautti and Lau consider to be the beginning of hemmolytiques conclusive demonstration of EICA, paving the way to additional experiments to identify the mechanisms underlying the evolution of the invasive populations of H.

Hemptinne conceived and designed the work. Facon revised the article critically. Ducamp performed the chemical analyses. Magro did the statistical analyses.

Grenouillet for statistical advice and G. Garrigues hemolytoques technical support. The evolution of chemical defenses along invasion routes: Harmonia axyridis Pallas Coccinellidae: Coleoptera as a case study.

National Center for Biotechnology InformationU. Journal List Ecol Evol v. Ansmies online Jul Author information Article notes Copyright and License information Disclaimer. This is an open access article under the terms of the http: Extraction anemeis harmonine from the eggs Following the extraction of the ECHCs, 48 to 50 eggs per population were analyzed for harmonine content.

Comparison of the ECHCs The average mass of the eggs was calculated on samples of 48 to 50 eggs for each of the three populations. Comparison of harmonine concentration in the eggs The concentration of harmonine in the eggs laid by the ladybirds of the three populations was analyzed using a GLM with a Poisson family.

Open in a separate window. Kovats retention hemolytiwues NI: Biometrics62 1— Evolution66— Methods in chemical ecology. NeoBiota107— Developmental and Comparative Immunology49— Elytra color as a signal of chemical defense in the Asian ladybird beetle Harmonia axyridis. Behavioral Ecology and Sociobiology61— A new colony structure of the invasive Argentine ant Linepithema hemolytiqies in Southern Europe.

Biological Invasions12—