# The spider mite Oligonychus yothersi (Acari: Tetranychidae), a new alien pest in Europe: joint records from insular Portugal, Spain and Italy revealed by integrative taxonomy

Lurdes Borges Silva, Alain Migeon, Philippe Auger, Francesco Andolina, Francisco Ferragut, Thomas Giordano, Pedro Naves, Luis Silva, Haralabos Tsolakis, Denise Navia

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Lurdes Borges Silva, Alain Migeon, Philippe Auger, Francesco Andolina, Francisco Ferragut, et al. The spider mite Oligonychus yothersi (Acari: Tetranychidae), a new alien pest in Europe: joint records from insular Portugal, Spain and Italy revealed by integrative taxonomy. Acarologia, 2025, 65 (4), pp.1213-1224. 10.24349/vjai-u7f7. hal-05484077

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# The spider mite *Oligonychus yothersi* (Acari: Tetranychidae), a new alien pest in Europe: joint records from insular Portugal, Spain and Italy revealed by integrative taxonomy

Lurdes Borges Silva IDa,b,c,d, Alain Migeon IDa, Philippe Auger IDa, Francesco Andolina Id,e, Francisco Ferragut If, Thomas Giordano Id,e, Pedro Naves Id,g, Luis Silva Id,b,c,d, Haralabos Tsolakis Id,e, Denise Navia Id,a

## Original research

### ABSTRACT

The spider mite *Oligonychus yothersi* (McGregor, 1914) is for the first time reported in Europe from four European insular regions with mild oceanic and Mediterranean climates- the Azores and Madeira archipelagos (Portugal), Canary Islands (Spain) and Sicily (Italy). Although populations were detected in the Canary and Azores islands since 2006 and 2015, respectively, their taxonomic identification remained unresolved due to the lack of male specimens and the unavailability of DNA reference sequences. An integrative taxonomic approach based on morphological traits of two males collected in September 2024 and on recently and previously obtained COI DNA sequences from multiple localities and host plants samples allowed its accurate identification and host plant records. Specimens have been collected from a range of 21 host plants, including native, cultivated and invasive species. New host plant records suggest host adaptation in the colonized areas. Six COI haplotypes have been identified in the studied areas. COI DNA sequences obtained from accurately identified specimens which can support further identifications are provided. Its pest status on other continents is detailed along with the warning on the risk that this spider mite poses for Europe.

**Keywords** Mangifera indica; Vitis vinifera; Tetranychidae; invasive phytophagous mites; host adaptation; plant protection

### Introduction

Arthropod invasions represent a growing concern both worldwide (Venette and Hutchison, 2021) and at the regional level, particularly in insular ecosystems (Cardoso et al. 2009; Florencio et al. 2013; Borges et al. 2013). Spider mites (Tetranychidae) include some of the most important agricultural pests, raising increasing concern due to their i) broad distribution

Received 08 October 2025
Accepted 25 November 2025
Published 04 December 2025

Corresponding author
Denise Navia ID:
denise.navia@inrae.fr

Academic editor
Kreiter, Serge

https://doi.org/10.24349/vjai-u7f7

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

<figure><img src="image_2.png" /></figure>

Borges Silva L. et al.

Licensed under
Creative Commons CC-BY 4.0

OPEN ACCESS

<footnote>a CBGP, INRAE, CIRAD, Institut Agro, IRD, Univ Montpellier, Montpellier, France.</footnote>

<footnote>b CIBIO, Research Centre in Biodiversity and Genetic Resources, University of the Azores, Ponta Delgada, São Miguel, Portugal.</footnote>

<footnote>c BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Vairão, Portugal.</footnote>

<footnote>d UNESCO Chair -Land Within Sea: Biodiversity and Sustainability in Atlantic Islands, Universidade dos Açores, R. Mãe de Deus 13A, 9500-321 Ponta Delgada, Portugal.</footnote>

<footnote>e Department of Agricultural, Food and Forest Sciences, University of Palermo, Palermo, Italy.</footnote>

<footnote>f Instituto Agroforestal Mediterráneo, Universitat Politècnica de València, Valencia, Spain.</footnote>

<footnote>g Instituto Nacional de Investigação Agrária e Veterinária, I.P. (INIAV), Oeiras, Portugal.</footnote>
and host range (Migeon and Dorkeld, 2025); ii) ability to rapidly colonize new areas and adapt to new host plants (Boubou *et al.* 2011); iii) populations can be favored by host plants under water stress (Migeon *et al.* 2023); and iv) economic impact on agricultural, ornamental and forest crops (Hoy 2011).

Along the last decades some alien spider mites became invasive pests in Europe causing impact on agricultural crops (Navajas *et al.* 2010; Auger *et al.* 2023; Ferragut *et al.* 2013). Remarquable examples are *Oligonychus perseae* Hirst on avocado, and *Eutetranychus orientalis* (Klein) and *E. banksi* (McGregor) on citrus, for which control remains challenging (Torres *et al.* 2023; López-Olmos and Ferragut 2023).

The most effective way to manage the risks posed by invasive pests is to prevent their establishment in new areas. Early detection has critical role in this process, enabling rapid response measures before pests can spread and cause impacts (EPPO 2021; Desneux *et al.* 2022). Carrying out an accurate identification is essential for surveillance, early warning systems, and the implementation of targeted management strategies. However, for certain groups of organisms—such as phytophagous mites—early detection is particularly challenging, due to their cryptic habitus and absence of symptoms at low population densities (Dhooria 2016; Vargas Madriz *et al.* 2025). Moreover, taxonomic identification can be particularly challenging due to the limited number of reliable morphological diagnostic traits (Ovalle *et al.* 2020; Razuvaeva *et al.* 2023), absence or low number of male specimens for some species, and the lack of available genetic data, even for groups of significant agricultural importance such as spider mites (Navia *et al.* 2011; Migeon and Dorkeld 2025).

Herein we report the first occurrence of *Oligonychus yothersi* (McGregor, 1914) in Europe, which was detected from four insular territories – the Macaronesian Azores, Madeira (Portugal) and Canary (Spain) archipelagos, and the largest Mediterranean island, Sicily (Italy). Host plants in the studied areas are presented, including three new host families – Apocynaceae, Myricaceae, Pittosporaceae. Haplotypes have been characterized.

# Materials and methods

## Mite collection

In the Azores, Portugal, surveys of plant inhabiting mites were carried out on exotic (both cultivated or invasive) and native plant species in July 2015 and in May and September 2024, in São Miguel. Faial, Flores and Pico islands by AM, DN, FF, LBS, LS, PA. In Madeira, surveys were carried out in 2022 by PA, AM, PN and DN. In Palermo and Messina provinces, Sicily, Italy, in April and July 2025 on mango trees by FA, TG, HT and Gabriela Lo Verde. We have also added an older sample collected by FF, AM and Maria Navajas in the Canary Islands in 2006. Collection localities and data are detailed in Table 1. Specimens were collected directly from symptomatic rusting or chlorotic leaves with a brush under a stereomicroscope. Specimens were preserved in 100% ethanol for molecular analysis, and in 70% ethanol, for morphological studies.

## Morphological identification

Mites were cleared in lactic acid (50%) for a day and mounted in Hoyer's medium for phase contrast and differential interference contrast (DIC) microscope observation. Morphological identification was performed using the key of Mushtaq *et al.* (2021) for the genus *Oligonychus* as well as the descriptions of *O. yothersi* by McGregor (1914, 1950) and Pritchard and Baker (1995) and of closely related species. Voucher specimens are deposited at the Centre de Biologie pour la Gestion des Population, 2018, "CBGP - Continental Arthropod Collection", Montferrier-sur-Lez, France, https://doi.org/10.15454/D6XAKL, in Zoological Collection of Arthropods at CIBIO-Açores, Ponta Delgada, São Miguel, Portugal and in Acarological Collection at SAAF Department, Palermo University, Italy.
<table><caption>**Table 1** *Oligomychus yothersi* collection data in Europe, including host plants/families, collection sites and dates. Samples marked with an asterisk (*) were used for morphological analyses. New host plant records are in bold.</caption><thead><tr><th>Sample name</th><th>Country</th><th>Island</th><th>Elementary administrative unit</th><th>Locality</th><th>Date of collect</th><th>Latitude</th><th>Longitude</th><th>Plant family</th><th>Plant species</th></tr></thead><tbody><tr><td>2006-095</td><td>Canary Is.</td><td>Tenerife</td><td>La Orotava</td><td>Plaza de la Constitución</td><td>2006-X-22</td><td>28.39346</td><td>-16.52189</td><td>Myrtaceae</td><td>Myrtus communis</td></tr><tr><td>2022-002</td><td>Madeira Is.</td><td>Madeira</td><td>Câmara de Lobos</td><td>Jesus Maria José</td><td>2022-V-17</td><td>32.66175</td><td>-16.98061</td><td>Anacardiaceae</td><td>Mangifera indica</td></tr><tr><td>2022-035</td><td>Madeira Is.</td><td>Madeira</td><td>Santana</td><td>E211 Faja Alta -São Jorge</td><td>2022-V-18</td><td>32.81748</td><td>-16.89864</td><td>Annonaceae</td><td>Annona cherinola</td></tr><tr><td>2022-043</td><td>Madeira Is.</td><td>Madeira</td><td>Santana</td><td>Arco de São Jorge, Museu do Vinho</td><td>2022-V-19</td><td>32.82431</td><td>-16.95375</td><td>Lauraceae</td><td>Laurus novocanariensis</td></tr><tr><td>2015-156</td><td>Azores Is.</td><td>Faial</td><td>Flamengos</td><td>Jardim botânico</td><td>2015-X-07</td><td>38.5508</td><td>-28.6392</td><td>Platanaceae</td><td>Platanus hispanica</td></tr><tr><td>2015-158</td><td>Azores Is.</td><td>Faial</td><td>Pedro Miguel</td><td>Praia da Almoxarife</td><td>2015-X-07</td><td>38.5677</td><td>-28.617</td><td>Theaceae</td><td>Camellia japonica</td></tr><tr><td>2015-162</td><td>Azores Is.</td><td>Faial</td><td>Castelo Branco</td><td></td><td>2015-X-08</td><td>38.5419</td><td>-28.7447</td><td>Rosaceae</td><td>Malus domestica</td></tr><tr><td>2015-173</td><td>Azores Is.</td><td>Pico</td><td>Terra do Pao</td><td>Lajes do Pico, Hotel Aldeia da Fonte</td><td>2015-X-06</td><td>38.4139</td><td>-28.2833</td><td>Solanaceae</td><td>Lycium europaeum</td></tr><tr><td>2024-214</td><td>Azores Is.</td><td>São Miguel</td><td>Sete Cidades</td><td>Freiras</td><td>2024-IX-17</td><td>37.85838</td><td>-25.78351</td><td>Pittosporaceae</td><td>Pittosporum undulatum</td></tr><tr><td>2024-217</td><td>Azores Is.</td><td>São Miguel</td><td>Sete Cidades</td><td>Freiras</td><td>2024-IX-17</td><td>37.85805</td><td>-25.78312</td><td>Fagaceae</td><td>Quercus robor</td></tr><tr><td>2024-229</td><td>Azores Is.</td><td>São Miguel</td><td>Sete Cidades</td><td>Freiras</td><td>2024-IX-17</td><td>37.86093</td><td>-25.78515</td><td>Myricaceae</td><td>Myrica faya</td></tr><tr><td>2024-240</td><td>Azores Is.</td><td>São Miguel</td><td>Sete Cidades</td><td>Lagoa Azul West</td><td>2024-IX-18</td><td>37.86892</td><td>-25.79477</td><td>Fabaceae</td><td>Acacia melanoxylon</td></tr><tr><td>2024-265</td><td>Azores Is.</td><td>São Miguel</td><td>Sete Cidades</td><td>Lagoa Azul West</td><td>2024-IX-18</td><td>37.86918</td><td>-25.79355</td><td>Polygonaceae</td><td>Reynoutria japonica</td></tr><tr><td>2024-267</td><td>Azores Is.</td><td>São Miguel</td><td>Sete Cidades</td><td>Freiras</td><td>2024-IX-17</td><td>37.86002</td><td>-25.78368</td><td>Fagaceae</td><td>Quercus palustris</td></tr><tr><td>2024-287</td><td>Azores Is.</td><td>São Miguel</td><td>Nordeste</td><td>C. de Praia do Lombo Gordo</td><td>2024-IX-20</td><td>37.7861</td><td>-25.145959</td><td>Platanaceae</td><td>Platanus hispanica</td></tr><tr><td>2024-290</td><td>Azores Is.</td><td>São Miguel</td><td>Nordeste</td><td>C. de Praia do Lombo Gordo</td><td>2024-IX-20</td><td>37.7861</td><td>-25.145959</td><td>Lauraceae</td><td>Persea indica</td></tr><tr><td>2024-299</td><td>Azores Is.</td><td>São Miguel</td><td>Maia</td><td>Cha Gorreana</td><td>2024-IX-20</td><td>37.81885</td><td>-25.40184</td><td>Theaceae</td><td>Camellia japonica</td></tr><tr><td>2024-302</td><td>Azores Is.</td><td>São Miguel</td><td>Nordeste</td><td>Miradouro da Ponta da Madrugada</td><td>2024-IX-20</td><td>37.78921</td><td>-25.14609</td><td>Rosaceae</td><td>Rosa sp.</td></tr><tr><td>2024-310</td><td>Azores Is.</td><td>São Miguel</td><td>Maia</td><td>Cha Gorreana</td><td>2024-IX-20</td><td>37.81885</td><td>-25.40184</td><td>Fagaceae</td><td>Castanea sativa</td></tr><tr><td>2024-318</td><td>Azores Is.</td><td>São Miguel</td><td>Nordeste</td><td>C. de Praia do Lombo Gordo</td><td>2024-IX-20</td><td>37.78516</td><td>-25.14575</td><td>Vitaceae</td><td>Vitis sp.</td></tr><tr><td>2024-323</td><td>Azores Is.</td><td>São Miguel</td><td>Rosto Do Cão (Livramento)</td><td>Rua do Cacto</td><td>2024-IX-21</td><td>37.75047</td><td>-25.6108</td><td>Apocynaceae</td><td>Plumeria alba</td></tr><tr><td>2024-330</td><td>Azores Is.</td><td>Pico</td><td>São Caetano</td><td>Ponta des Formigas</td><td>2024-IX-22</td><td>38.42194</td><td>-28.42837</td><td>Pittosporaceae</td><td>Pittosporum undulatum</td></tr><tr><td>2024-352</td><td>Azores Is.</td><td>Pico</td><td>São Caetano</td><td>Ponta des Formigas</td><td>2024-IX-22</td><td>38.42194</td><td>-28.42837</td><td>Vitaceae</td><td>Vitis vinifera</td></tr><tr><td>2024-358</td><td>Azores Is.</td><td>Pico</td><td>São João</td><td>Parque Florestal e Recreio de São João Pequenino</td><td>2024-IX-23</td><td>38.41794</td><td>-28.36073</td><td>Fabaceae</td><td>Acacia melanoxylon</td></tr><tr><td>2024-366</td><td>Azores Is.</td><td>Pico</td><td>São João</td><td>Parque Florestal e Recreio de São João Pequenino</td><td>2024-IX-23</td><td>38.41794</td><td>-28.36073</td><td>Myricaceae</td><td>Myrica faya</td></tr><tr><td>2024-369</td><td>Azores Is.</td><td>Pico</td><td>São João</td><td>Parque Florestal e Recreio de São João Pequenino</td><td>2024-IX-23</td><td>38.41794</td><td>-28.36073</td><td>Platanaceae</td><td>Platanus hispanica</td></tr><tr><td>2024-381</td><td>Azores Is.</td><td>Pico</td><td>Madalena</td><td>EN3 Dog shelter</td><td>2024-IX-24</td><td>38.51964</td><td>-28.50095</td><td>Lauraceae</td><td>Persea indica</td></tr><tr><td>2024-397</td><td>Azores Is.</td><td>Pico</td><td>São Caetano</td><td>Ponta des Formigas</td><td>2024-IX-22</td><td>38.42194</td><td>-28.42837</td><td>Fabaceae</td><td>Acacia melanoxylon</td></tr><tr><td>2024-405</td><td>Azores Is.</td><td>Flores</td><td>Santa Cruz das Flores</td><td>Ribeira dos Barqueiros</td><td>2024-IX-27</td><td>39.44961</td><td>-31.1398</td><td>Fagaceae</td><td>Castanea sativa</td></tr><tr><td>2024-407</td><td>Azores Is.</td><td>Flores</td><td>Fajã Grande</td><td>Ponta</td><td>2024-IX-27</td><td>39.46965</td><td>-31.25458</td><td>Ericaceae</td><td>Rhododendron indicum</td></tr><tr><td>2024-427</td><td>Azores Is.</td><td>Flores</td><td>Fajã Grande</td><td>Ponta</td><td>2024-IX-27</td><td>39.46965</td><td>-31.25458</td><td>Platanaceae</td><td>Platanus hispanica</td></tr><tr><td>2025-250</td><td>Italy</td><td>Sicilia</td><td>Palermo</td><td>Via Luparello</td><td>2025-IX-07</td><td>38.1143</td><td>13.3005</td><td>Anacardiaceae</td><td>Mangifera indica</td></tr><tr><td>Italy</td><td>Sicilia</td><td>Messina</td><td>Messina</td><td>Milazzo</td><td>2025-VII-20</td><td>38.1134</td><td>15.1422</td><td>Anacardiaceae</td><td>Mangifera indica</td></tr><tr><td>Italy</td><td>Sicilia</td><td>Messina</td><td>Messina</td><td>Milazzo</td><td>2025-VII-20</td><td>38.114</td><td>15.1353</td><td>Anacardiaceae</td><td>Mangifera indica</td></tr></tbody></table>
## Molecular analysis

Total DNA from 64 isolated specimens, obtained from 34 samples (Table 1), was extracted using the DNeasy Blood & Tissue kit (Qiagen, Hilden, Germany), according to the DNA extraction protocol "Purification of Total DNA from Animal Blood or Cells" (Spin-Column Protocol) adapted for extracting total DNA from mites (Mendonça *et al.* 2011). The isolated DNA was directly used as a template for PCR amplification of two fragments of the mitochondrial cytochrome c oxidase subunit I (COI) gene using two primer sets: LCO1490 (5'-GGTCAACAAATCATAAAGATATTGG-3') and HCO2198 (5'-TAAACTTCAGGGTGACCAAAAAATCA-3') (Folmer *et al.* 1994); C1J1718 (5'-GGAGGATTTGGAAATTGATTAGTTCC-3') and COIrevA (5'-CCWGTYARMCCHCCWAY-AGTAAA-3') (Auger *et al.* 2023). For a final volume of 25 µL, the PCR mixture consisted of 12.5 µL of Qiagen Multiplex PCR Master Mix (2X), 1.25 µL of each primer (10 µM), 6 µL of nuclease-free water, and 4 µL of template DNA. The PCR conditions for LCO1490/HCO2198 were conducted under the following thermal cycling conditions: an initial denaturation at 95 °C for 15 min; followed by 40 cycles of denaturation at 95 °C for 45 s, hybridation at 45 °C for 30 s, and elongation at 72 °C for 70 s; with a final elongation at 72 °C for 10 min. Amplification with C1J1718/COIrevA was performed with an initial denaturation at 94 °C for 15 min, followed by 5 cycles of denaturation at 94 °C for 60 s, hybridation at 40 °C for 90 s, and elongation at 62 °C for 90 s; then 30 additional cycles of denaturation at 94 °C for 60 s, hybridation at 45 °C for 90 s, and elongation at 62 °C for 90 s; and a final elongation at 62 °C for 10 min. The amplified PCR products were sequenced by Eurofins Genomics Europe Pharma and Diagnostics (Sanger/PCR GmbH, Cologne Branch, Germany), using the same primers as in the PCR amplification.

Sequences were edited using Codon Aligner v.4.1.1 (CodonCode Corporation) and aligned using the default parameters. All the sequences were verified for stop codons and for insertions/deletions.

When it was possible, the two overlapping sequences were concatenated for phylogenetic analysis. New sequences were combined with all *Oligonychus* s. str. COI sequences longer than 400 pb retrieved from GenBank (one sequence per species, see Supplementary file S2). Six *Panonychus* species, were used as outgroup as proposed by Matsuda *et al.* (2018). The dataset obtained contained 28 sequences of 1268 pb (when aligned). A maximum likelihood (ML) analysis was conducted using Mega 12 (v12.0.15) (Kumar *et al.* 2024) with G+I model and fast adaptive bootstrap (Supplementary file S3).

## Results and discussion

The red mite *O. yothersi* (Figure 1) was identified from four European insular regions: in the Macaronesia (Atlantic), the Azores and Madeira archipelagos (Portugal), the Canary Islands (Spain); and the Sicily Island (Italy) in the central Mediterranean sea.

Identification was based on an integrative taxonomic approach. Since male specimens are required for species-level identification in the *Oligonychus* genus, morphological identification was only possible for two of the 26 samples collected in the Azores in September 2024, which were the only ones containing each one male specimen. Perfect laterally mountings of these males enabled the reliable morphological taxonomic identification at species level. No *O. yothersi* DNA reference sequences being available in public databases, COI DNA sequences obtained from females of the morphologically identified two populations were then used as reference and enabled identification of all other populations. The delayed identification of this invasive mite in Europe — nearly twenty years after the first symptom observations — is mainly due to the species thelytokus parthenogenetic reproduction which can result in the absence of males (Ferla *et al.* 2024) and the lack of barcoding DNA sequences, highlighting the importance of availability of reference sequences in databases to support early detection and monitoring of invasive pests.
<figure><img src="image_3.png" /><figcaption>**Figure 1** *Oligonychus yothersi* living specimens on *Vitis vinifera*: A – Two adult females; B – Nymphs and eggs.</figcaption></figure>

**Family Tetranychidae Donnadieu, 1875**

**Tribe Tetranychini Reck, 1950**

**Genus *Oligonychus* Berlese, 1886**

Type species *Heteronychus brevipodus* Targioni -Tozzetti, 1878: 255

**Oligonychus (Oligonychus) yothersi (McGregor, 1914)**

**Morphological identification**

Distinctive characters (Figure 2): females of this species bear 7 tactile setae on the tibia of the leg I and have the first pair of dorsocentral setae ($c_1$) longer than the interval between the setae $c_1$ and $d_1$ (Figure 2 A-C), belonging to the *coffeae* species subgroup (Mushtaq et al. 2021). In this subgroup of 44 species, the male aedeagus of *O. yothersi* is unique, with the shaft deflexed sharply more than 90° from the axis and forming a hooked portion that is longer than the shaft, tapering to a thin (unbarbed) truncate tip (McGregor, 1950). According to McGregor's (1950) drawing of the aedeagus of *O. yothersi*, there is an indentation on the proximal part of the dorsal margin, which was also observed in both males collected (Figure 2 D, E). In addition, in several female specimens, we observed additional solenidia on the tarsus (from one to four) and tibia (from one to three) of the leg I as reported by Pritchard and Baker (1955).

**Molecular analysis**

Among the 34 samples collected, 31 gathering 1 to 4 individuals per sample, were successfully sequenced for the COI-C1J1718 fragment, yielding to 63 sequences of 876 bp, and 23, each gathering 1 to 4 individuals per sample, were successfully sequenced for the COI-LCO-HCO fragment, yielding to 48 sequences of 672 bp. The two fragments overlap by 462 pb and the variable nucleotide sites were present only on the COI-C1J1718 fragment. These sites and the Genbank sequence identifiers are reported in Table 2.

Interestingly, all the sequences showed high similarity, with 49 individuals sharing same haplotype, only 11 individuals sharing one mutation and four individuals with other mutations on the COI-C1J1718 (Table 2, Supplementary table 1). No genetic structuration was observed in regard of the geographic origin nor the host plants. Notably, the most common haplotype was shared by the majority of Azoreans and Madeiran specimens and all the four Sicilian specimens. The phylogenetic tree obtained is shown in Figure 3.
<figure><img src="image_4.png" /><figcaption>**Figure 2** *Oligonychus yothersi* female: A–B – 7 tactile setae and one solenidion on tibia I (A – dorsal view, B – ventral view); C – dorsohys-terosomal setae *c*₁ longer than the distance *c*₁–*d*₁; male: D – aedeagus of the male collected on *Vitis* sp., E – aedeagus of the male collected on *Acacia melanoxylon*. Scale bars A–C = 20µm, D–E = 10µm.</figcaption></figure>

Distribution

*Oligonychus yothersi* was described from the United States in 1914 (McGregor), being
currently widespread (Figure 4) in the Americas (from the United States to Argentina and

<table><caption>**Table 2** Variable nucleotide sites in mitochondrial *COI* sequences resulting in six *COI* haplotypes (H1–H6) and Genbank accession identifiers.</caption><thead><tr><th rowspan="2">Haplotypes</th><th colspan="5">Variables sites</th><th rowspan="2">Count</th><th colspan="2">Genbank accession Ids</th></tr><tr><th>C1J1718 position</th><th>312</th><th>361</th><th>409</th><th>579</th><th>801</th><th>C1J1718</th><th>LCO</th></tr></thead><tbody><tr><td>LCO position</td><td>516</td><td>565</td><td>613</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>H1</td><td>C</td><td>G</td><td>G</td><td>A</td><td>G</td><td>49</td><td>PX353980</td><td>PX353976</td></tr><tr><td>H2</td><td>.</td><td>.</td><td>.</td><td>G</td><td>.</td><td>10</td><td>PX353981</td><td></td></tr><tr><td>H3</td><td>A</td><td>.</td><td>.</td><td>.</td><td>.</td><td>1</td><td>PX353982</td><td>PX353977</td></tr><tr><td>H4</td><td>.</td><td>.</td><td>A</td><td>.</td><td>.</td><td>1</td><td>PX353983</td><td>PX353978</td></tr><tr><td>H5</td><td>.</td><td>.</td><td>.</td><td>.</td><td>A</td><td>1</td><td>PX353984</td><td></td></tr><tr><td>H6</td><td>.</td><td>T</td><td>.</td><td>G</td><td>.</td><td>1</td><td>PX353985</td><td>PX353979</td></tr></tbody></table>
<figure><img src="image_5.png" /><figcaption>**Figure 3** Maximum likelihood (ML) phylogenetic tree based on the mitochondrial *COI* gene using the G+T Gamma model. Bootstrap values (>50%) based on adaptive bootstrap (Mega 12) are indicated at nodes. Genbank accession numbers are indicated.</figcaption></figure>

Chile), and has also been recorded in Asia (China) (Ma and Yuan, 1981) and the Middle East (Iran) (Sheikholeslam-Zadeh and Sadeghi-Nameghi, 2010). No prior records were known from Europe (Migeon and Dorkeld 2025). The record from Hawaii has never been confirmed, suggesting that the species may not be present in the Hawaiian Islands (Goff, 1986).

In this study, *O. yothersi* was collected from four of the nine Azorean islands: Faial, Flores, Pico, and São Miguel, including eastern, central and western island-groups. These results show that the mite is widespread in the Azorean archipelago, and can possibly occur on other Azorean islands where similar symptoms have been observed. In the Macaronesia, besides Azores, the mite was identified from three samples from Madeira and from one sample from Tenerife (Canary Islands). In addition, the species was identified from Sicily in the central Mediterranean Sea. Up to date the mite was not recorded from continental Europe, but further surveys should be conducted to assess its eventual distribution on the continent and to define measures to prevent its dissemination. Considering the new insular detections in Europe, the updated distribution map of *O. yothersi* is presented in Figure 4.

**Introduction history and pathways**

The wide distribution of *O. yothersi* in the Macaronesia area, as well as the high number of new host plants (Table 1), including native ones, suggest that its introduction in the region is not a recent event. Furthermore, its detection in samples collected in 2006 suggests an introduction
<figure><img src="image_6.png" /><figcaption>**Figure 4** World distribution map of *Oligonychus yothersi*. TDWG 4 (TDWG, 2025) level and known occurrences derived from GBIF (2025) and Migeon and Dorkeld, (2025). New records from this study.</figcaption></figure>

several years distant. The absence of highly conspicuous symptoms on plants of economic importance can explain how the species remained undetected for decades in Macaronesia, while in Sicily a high level of infestation with damages on cultivated plants was obvious. In the absence of significant genetic variability among populations, it would be interesting to determine the ecological and environmental conditions that explain why this mite is causing damage only on Sicily, and not in the Macaronesian Archipelagos.

The homogeneity of all the sequences suggests a single introduction. The high level of symptoms in Sicily suggests that the introduction, or at least the pest emergence, is recent and could not fail to be noticed. This homogeneity and the lack of reference sequences from the native area does not allow drawing more hypothesis on invasion pathways and history. However, based on evidence from other invasive spider mites, such as *Tetranychus evansi* Baker & Pritchard, 1960, *Eotetranychus lewisi* (McGregor, 1943) and *O. perseae*, which share comparable bioecological traits, plant trade is considered the most probable pathway of introduction (Navajas *et al.* 2013; EFSA 2021; Torres *et al.* 2024).

## Host plant symptoms, pest status and potential risks for Europe

The spider mite *O. yothersi* is a highly polyphagous species recorded on more than 80 host plant species belonging to over 40 botanical families. Plant families most commonly reported as *O. yothersi* hosts are Rosaceae, Myrtaceae and Fabaceae (Migeon and Dorkeld, 2025).

In this study, *O. yothersi* was collected on 21 plant species belonging to 16 angiosperm families (Table 1). Among them, there are native and Macaronesia endemic species such as *Laurus novocanariensis* Rivas Mart., Lousã, Fern.Prieto, E.Días, J.C.Costa & C.Aguiar, and *Myrica faya* Dryand., invasive species such as *Acacia melanoxylon* R. Br. and *Pittosporum undulatum* Vent.; and agricultural, forestry or ornamental species such as *Annona cherimola* Mill., *Fagus sylvatica* L., *Malus domestica* Borkh., *Mangifera indica* L., *Myrtus communis* L., *Persea indica* (L.) Spreng., *Platanus hispanica* Mill. ex Münchh., *Plumeria alba* L., *Quercus palustris* Münchh., *Quercus robur* L., *Castanea sativa* Mill., *Rosa* sp., *Reynoutria japonica* Houtt. and *Vitis vinifera* L. The high number of host plants in the Macaronesia, including several new reports (families and species) and endemic plants, suggests that the red mite has
<figure><img src="image_7.png" /><figcaption>**Figure 5** Feeding damage caused by *Oligonychus yothersi* in the Azores Archipelago and Sicily (Italy). A – Yellowing and necrosis on *Persea indica* (Azores). B – Central leaf bronzing on *Camellia japonica* (Azores). C – Typical feeding symptoms on *Mangifera indica* (Sicily). D – Colony of *O. yothersi* with two *Stethorus* sp. larvae (predators) on *Mangifera indica* (Sicily).</figcaption></figure>

adapted to new hosts along the establishment process, and confirm its polyphagy as reported from other continents. It is worth noting that *O. yothersi* was found infesting *P. undulatum*, one of the most aggressive and widespread invasive plants in the Azorean archipelago (Borges Silva *et al.* 2018, 2022). In Sicily and Madeira, *O. yothersi* was collected from mango orchards, as it also has been reported from several countries in Central and South America (see Ochoa *et al.* 1990, and for the last update Migeon and Dorkeld, 2025). Mites were observed primarily on the upper surface of the leaves, but colonies were also found on the undersides, sometimes in association with *Stethorus* sp. larvae(Coleoptera: Coccinellidae), which are known predators of tetranychid mites (Figure 5).

*Oligonychus yothersi* has been reported as a major foliar pest of avocado (*Persea americana* Mill.) under the name of ‘avocado red mite’ in Central and South America (Rioja *et al.* 2019), and it has also been reported causing damages in ‘erva-mate’, *Ilex paraguariensis* A.ST.-Hil. (Aquifoliaceae) in South Brazil, (Dameda *et al.* 2021; Rode *et al.* 2023) and recently on vine (*Vitis vinifera* L.) in Brazil (Ferla *et al.*, 2024).

Considering the current host plant range of *O. yothersi* around the world (Migeon and Dorkeld, 2025) the main concerns in Europe would be for the mango, avocado and vine plantations in Mediterranean areas such as Italy, France, Portugal and Spain. Other important crops, such as *Prunus* and *Malus* fruit trees, are also widely cultivated in continental Europe and could potentially be affected. Additionally, this mite could also cause some impact to the recent
development of tea plantations in continental Europe (Tea Grown in Europe Association, 2023) Its association with ornamental plants can contribute to its dissemination across the European continent, and facilitate its establishment in new areas a process that would also be reinforced by its association with other plant species, such as native oaks (*Quercus* spp.) and others. As a precautionary measure, phytosanitary measures should be implemented at a regional level to avoid its spread to continental areas and minimize potential impacts.

## Acknowledgements

Main funding was provided by the project "AllInterAz - Close encounters in Atlantic islands-unravelling arthropod-alien plant multitrophic interactions in the Azores islands" (BIOPRECH-RZ110-ALINTERAZ/BIOPOLIS). Sequencing cost was supported by PrepAcari project (Petits mais Costauds : de la taxonomie intégrative à l'écologie pour se préparer au risque acarien) IB SPE 2023 INRAE. Lurdes Borges Silva was also supported by National Funds through FCT- Fundação para a Ciência e a Tecnologia in the scope of the project UID/50027 – Rede de Investigação em Biodiversidade e Biologia Evolutiva. Pedro Naves would like to acknowledge funding provided by the Fundação para Ciência e Tecnologia (FCT/Portugal) project EXPL/ASP-AGR/0082/2021, and the “GREEN-IT Bioresources for Sustainability” https://doi.org/10.54499/UIDB/04551/2020. Surveys carried out in Sicily were funded by the University of Palermo (Fondo di Finanziamento per la Ricerca FFR, by Gabriella Lo Verde and Haralabos Tsolakis).

## ORCID

Lurdes Borges Silva https://orcid.org/0000-0002-4185-733X

Alain Migeon https://orcid.org/0000-0003-4041-5158

Philippe Auger https://orcid.org/0000-0002-2671-9561

Francesco Andolina https://orcid.org/0009-0001-1314-8739

Francisco Ferragut https://orcid.org/0000-0003-1545-6011

Thomas Giordano https://orcid.org/0009-0003-3209-3192

Pedro Naves https://orcid.org/0000-0002-8297-5527

Luis Silva https://orcid.org/0000-0002-3434-6056

Haralabos Tsolakis https://orcid.org/0000-0001-8018-9496

Denise Navia https://orcid.org/0000-0003-3716-4984

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