# Contrasting Invasion Strategies, Convergent Outcomes: Establishment of Zaprionus tuberculatus and Ceroplastes ceriferus in Italy

**Authors:** Francesco Nugnes, Carmela Carbone, Fortuna Miele, Feliciana Pica, Sara Pierro, Raffaele Sasso, Mariagrazia Bodini, Umberto Bernardo

PMC · DOI: 10.3390/insects17020198 · Insects · 2026-02-12

## TL;DR

Two invasive insect species, a fruit fly and a wax scale, have successfully established in Italy despite limited genetic diversity, showing how ecological flexibility and human activity can aid invasions.

## Contribution

The study reveals how contrasting invasion strategies can lead to successful establishment despite low genetic variation, emphasizing the role of human-mediated transport and ecological plasticity.

## Key findings

- Zaprionus tuberculatus was found on ten host plants, with highest abundance on persimmon, and reproduced only on figs and blueberries.
- Ceroplastes ceriferus showed polyphagy and established on four host plants in southern Italy.
- Both species had limited genetic variation in Italy, yet successfully established, highlighting the genetic paradox in invasive species.

## Abstract

Two non-native insect species, the fruit fly Zaprionus tuberculatus and the wax scale Ceroplastes ceriferus, have recently extended their distribution range into Italy. Although they differ greatly in behaviour and biology, the fruit fly being highly mobile and the wax scale remaining fixed on the plant, both established themselves in local environments through different invasion pathways. This study examined where these species occur, which plants they attack, and whether they are able to reproduce under Italian conditions. The insects were also analysed genetically to understand how much variation they possess and how Italian populations relate to those from other parts of the world. Such information helps clarify how these species arrive, spread, and adapt to new areas. The findings show that both insects are already established on several host plants and can persist under Mediterranean climate conditions. Despite carrying only limited genetic variation, both species successfully settled in the region. This outcome reflects what is known as the “genetic paradox”, in which invasive species can thrive even with reduced genetic diversity, highlighting the need for continued surveillance and a better understanding of the mechanisms that enable successful establishment.

Global warming and international trade are accelerating biological invasions, making the Mediterranean basin a hotspot for alien arthropods. Two invasive species, the drosophilid Zaprionus tuberculatus and the wax scale Ceroplastes ceriferus, were recently detected in central-southern Italy. Surveys conducted between 2023 and 2024 detected Z. tuberculatus across several sites in Campania and Lazio, and C. ceriferus in Campania. Zaprionus tuberculatus was reared from ten host plants, including three new records, and reached its highest abundance on persimmon. Laboratory assays showed that Z. tuberculatus adults reproduced only on intact fig fruits and blueberry, indicating a limited risk for most crops. Ceroplastes ceriferus was found on four hosts, confirming its polyphagy and establishment in southern Italy. Molecular analyses revealed two COI haplotypes in the Italian populations of both species, while three haplotypes were detected globally in Z. tuberculatus and seven in C. ceriferus. Sequence divergences were moderate, indicating limited but appreciable mitochondrial differentiation among geographic populations. In Z. tuberculatus, the COII haplotype detected in Italy did not match available African sequences, underscoring gaps in reference data and complicating the reconstruction of invasion routes. Despite contrasting ecological traits and dispersal strategies, both species achieved successful establishment in Italy, showing that ecological plasticity and human-mediated transport can offset reduced genetic diversity. These records expand the known European range of both species and highlight the need for coordinated surveillance, genetic monitoring, and preventive measures to limit further introductions in the Mediterranean region.

## Linked entities

- **Species:** Zaprionus tuberculatus (taxon 7297), Ceroplastes ceriferus (taxon 1028181)

## Full-text entities

- **Genes:** COX2 (cytochrome c oxidase subunit II) [NCBI Gene 4513] {aka COII, MTCO2}
- **Diseases:** Damage Fruits (MESH:D020263), injury to (MESH:D014947)
- **Chemicals:** ethanol (MESH:D000431), Deltamethrin (MESH:C017180), water (MESH:D014867), 2,6-di-tert-butyl-4-methylphenol (MESH:D002084), wax (MESH:D014885), acetoin (MESH:D000093), agarose (MESH:D012685), 3-(methylthio) propanol (MESH:C555618), GEA (-), ethyl acetate (MESH:C007650), methyl-eugenol (MESH:C005223)
- **Species:** Meleagris gallopavo (common turkey, species) [taxon 9103], Laurus nobilis (bay laurel, species) [taxon 85223], Vitis vinifera (wine grape, species) [taxon 29760], Morus sp. (in: birds) (species) [taxon 2047046], Coptodisca lucifluella (species) [taxon 1454946], Drosophila suzukii (species) [taxon 28584], Mangifera indica (mango, species) [taxon 29780], Annona cherimola (cherimoya, species) [taxon 49314], Fragaria x ananassa (strawberry, species) [taxon 3747], Photinia serratifolia (species) [taxon 140999], Tephritidae (fruit flies, family) [taxon 7211], Musa acuminata (banana, species) [taxon 4641], Carpophilus truncatus (species) [taxon 2944859], Eriobotrya japonica (loquat, species) [taxon 32224], Ficus carica (common fig, species) [taxon 3494], Zaprionus indianus (species) [taxon 76712], Drosophilidae (pomace flies, family) [taxon 7214], Diptera (flies, order) [taxon 7147], Vaccinium corymbosum (American blueberry, species) [taxon 69266], Persea americana (avocado, species) [taxon 3435], Drosophila melanogaster (fruit fly, species) [taxon 7227], Ficus microcarpa (Chinese banyan, species) [taxon 66385], Bactrocera dorsalis (oriental fruit fly, species) [taxon 27457], Juglans regia (English walnut, species) [taxon 51240], Diospyros kaki (Japanese persimmon, species) [taxon 35925], Leptocybe invasa (species) [taxon 1028344], Coccoidea (scale insects, superfamily) [taxon 33381], Ceratitis capitata (medfly, species) [taxon 7213], Zaprionus tuberculatus (species) [taxon 7297], Hymenoptera (hymenopterans, order) [taxon 7399], Ceroplastes japonicus (species) [taxon 1182649], Homo sapiens (human, species) [taxon 9606], Ribes rubrum (cultivated currant, species) [taxon 175228], Ziziphus jujuba (Chinese jujube, species) [taxon 326968], Tuta absoluta (species) [taxon 702717], Ceroplastes ceriferus (species) [taxon 1028181], Cameraria ohridella (species) [taxon 199129]
- **Mutations:** M165C

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## References

93 references — full list in the complete paper: https://tomesphere.com/paper/PMC12941184/full.md

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Source: https://tomesphere.com/paper/PMC12941184