Borges Silva, Lurdes ¹ ; Migeon, Alain ² ; Auger, Philippe ³ ; Andolina, Francesco ⁴ ; Ferragut, Francisco ⁵ ; Giordano, Thomas ⁶ ; Naves, Pedro ⁷ ; Silva, Luis ⁸ ; Tsolakis, Haralabos ⁹ and Navia, Denise ¹⁰

¹CBGP, INRAE, CIRAD, Institut Agro, IRD, Univ Montpellier, Montpellier, France & CIBIO, Research Centre in Biodiversity and Genetic Resources, University of the Azores, Ponta Delgada, São Miguel, Portugal & BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Vairão, Portugal & UNESCO Chair – Land Within Sea: Biodiversity and Sustainability in Atlantic Islands, Universidade dos Açores, R. Mãe de Deus 13A, 9500-321 Ponta Delgada, Portugal.
²CBGP, INRAE, CIRAD, Institut Agro, IRD, Univ Montpellier, Montpellier, France.
³CBGP, INRAE, CIRAD, Institut Agro, IRD, Univ Montpellier, Montpellier, France.
⁴Department of Agricultural, Food and Forest Sciences, University of Palermo, Palermo, Italy.
⁵Instituto Agroforestal Mediterráneo, Universitat Politècnica de València, Valencia, Spain.
⁶Department of Agricultural, Food and Forest Sciences, University of Palermo, Palermo, Italy.
⁷Instituto Nacional de Investigação Agrária e Veterinária, I.P. (INIAV), Oeiras, Portugal.
⁸CIBIO, Research Centre in Biodiversity and Genetic Resources, University of the Azores, Ponta Delgada, São Miguel, Portugal & BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Vairão, Portugal & UNESCO Chair – Land Within Sea: Biodiversity and Sustainability in Atlantic Islands, Universidade dos Açores, R. Mãe de Deus 13A, 9500-321 Ponta Delgada, Portugal.
⁹Department of Agricultural, Food and Forest Sciences, University of Palermo, Palermo, Italy.
¹⁰✉ CBGP, INRAE, CIRAD, Institut Agro, IRD, Univ Montpellier, Montpellier, France.

2025 - Volume: 65 Issue: 4 pages: 1213-1224

Original research

Keywords

Abstract

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 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 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.

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.

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 (c₁) longer than the interval between the setae c₁ and d₁ (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.

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 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 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 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 ''AlInterAz - 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).

acarologia_4846_supplementarytable1.xlsx

References

1. Auger P., Fossoud A., Migeon A. 2023. Three new alien spider mites (Prostigmata, Tetranychidae) from south-eastern France. Acarologia, 63(3): 826-833. https://doi.org/10.24349/yeys-kf03
2. Bacon S.J., Bacher S., Aebi A. 2012. Gaps in Border Controls Are Related to Quarantine Alien Insect Invasions in Europe. PLoS ONE, 7: e47689. https://doi.org/10.1371/journal.pone.0047689
3. Boieiro M., Varga-Szilay Z., Costa R., Crespo L., Leite A., Oliveira R., Pozsgai G., Rego C., Calado H., Teixeira M., Lopes D.H., Soares A., Borges P.A.V. 2024. New findings of terrestrial arthropods from the Azorean Islands. Biodiversity Data Journal 12: e136391. https://doi.org/10.3897/BDJ.12.e136391
4. Boller E.F. 1984. Eine einfache Ausschwemm-Methode zur schellen Erfassung von Raumilben, Trips und anderen Kleinathropoden im Weinbau. Schweiz Zeitschrift für Obst-und Weinbau, 120: 249-255.
5. Borges P.A.V., Reut M., Ponte N.B., Quartau J.A., Fletcher M., Sousa A.B., Pollet M., Soares A.O., Marcelino J., Rego C., Cardoso P. 2013. New records of exotic spiders and insects to the Azores, and new data on recently introduced species. Arquipélago Life and Marine Sciences, 30: 57-70.
6. Borges Silva L., Lourenço P., Teixeira A., Silva L. 2018. Biomass valorization in the management of woody plant invaders: The case of Pittosporum undulatum in the Azores. Biomass & Bioenergy, 109: 155-165. https://doi.org/10.1016/j.biombioe.2017.12.025
7. Borges Silva L., Pavão D.C., Elias R.B., Moura M., Ventura M.A., Silva L. 2022. Taxonomic, structural diversity and carbon stocks in a gradient of island forests. Scientific Reports, 12: 1038. https://doi.org/10.1038/s41598-022-05045-w
8. Borges Silva L., Madeira P., Pavão D.C., Elias R.B., Moura M., Silva L. 2023. Vascular plant taxa occurrences in exotic woodland and in natural and production forests on the Islands of São Miguel, Terceira, and Pico (Azores). Version 1.2. Universidade dos Açores. Sampling event dataset. Biodiversity Data Journal, 11: e109082. https://doi.org/10.3897/BDJ.11.e109082
9. Boubou A., Migeon A., Roderick G.K., Navajas M. 2011. Recent emergence and worldwide spread of the red tomato spider mite, Tetranychus evansi: genetic variation and multiple cryptic invasions. Biol. Invasions 13: 81-92. https://doi.org/10.1007/s10530-010-9791-y
10. Bragard C., Baptista P., Chatzivassiliou E., Di Serio F., Gonthier P., Jaques Miret J.A., Justesen A.F., Magnusson C.S., Milonas P., Navas-Cortes J.A., Parnell S., Potting R., Reignault P.L., Stefani E., Thulke H.-H., Van der Werf W., Vicent Civera A., Yuen J., Zappalà L., Grégoire J.-C., Malumphy C., Kertesz V., Maiorano A., MacLeod A. 2022. Scientific Opinion on the pest categorisation of Oligonychus perseae. EFSA Journal, 20: 7336. https://doi.org/10.2903/j.efsa.2022.7336
11. Brummitt R.K. 2001. World Geographic Scheme for Recording Plant Distributions, Edition 2. Hunt Institute for Botanical Documentation, Carnegie Mellon University (Pittsburgh). http://rs.tdwg.org/wgsrpd/doc/data/
12. Cardoso P., Lobo J.M., Aranda S.C., Dinis F., Gaspar C., Borges P.A.V. 2009. A spatial scale assessment of habitat effects on arthropod communities of an oceanic island. Acta Oecologica, 35: 590-597. https://doi.org/10.1016/j.actao.2009.05.005
13. Dameda C., Berté A.L.W., da Silva G.L., Johann L., Ferla N.J. 2021. Euseius concordis (Chant) (Acari: Phytoseiidae) as a potential agent for the control of yerba mate red mite Oligonychus yothersi (McGregor) (Acari: Tetranychidae). Phytoparasitica, 49(3): 377-383. https://doi.org/10.1007/s12600-020-00879-4
14. Desneux N., Blachere L., Arno J., Han P., Urbaneja A. 2022. Invasive pests in agroecosystems: detection, monitoring, and integrated management. Annual Review of Entomology, 67: 489-510.
15. Dhooria M.S. 2016. Fundamentals of Applied Acarology. Springer Nature, Singapore, 470 pp https://doi.org/10.1007/978-981-10-1594-6
16. EFSA Panel on Plant Health, Bragard C., Di Serio F., Gonthier P., Miret J.A.J., Justesen A.F., Magnusson C.S., Milonas P., Navas-Cortes J.A., Parnell S., Potting R., Reignault P.L., Thulke H.H., Van der Werf W., Civera A.V., Yuen J., Zappalà L., Gregoire J.C., Malumphy C., Czwienczek E., Kertesz V., Maiorano A., MacLeod A. 2021. Pest categorisation of Oligonychus mangiferus. Efsa Journal, 19(11): 6927. https://doi.org/10.2903/j.efsa.2021.6927
17. Ehara S., Gotoh T. 1992 Descriptions of two Panonychus spider mites from Japan, with a key to species of the genus in the world (Acari: Tetranychidae). Applied Entomology and Zoology, 27: 107-115. https://doi.org/10.1303/aez.27.107
18. Ehara S., Gotoh T. 1993 Description of the male of Panonychus thelytokus Ehara & Gotoh (Acari, Tetranychidae). Japanese Journal of Entomology, 61: 157-160.
19. EPPO. 2021. Guidelines on pest risk analysis and early detection. European and Mediterranean Plant Protection Organization. Available at: https://www.eppo.int
20. Ferla J.J., Granich J., Ferla N.J. 2024. Reproduction in the spider mite Oligonychus yothersi (Acari: Tetranychidae). Anais da Academia Brasileira de Ciencias, 96(4): e20230500.
21. Ferragut F., Navia D., Ochoa R. 2013. New mite invasions in citrus in the early years of the 21st century. Experimental and Applied Acarology, 59(1): 145-164. https://doi.org/10.1007/s10493-012-9635-9
22. Florencio M., Cardoso P., Lobo J.M., Azevedo E.B., Borges P.A.V. 2013. Arthropod assemblage homogenisation in oceanic islands: the role of exotic and indigenous species under landscape disturbance. Diversity and Distributions, 19: 1450-1460. https://doi.org/10.1111/ddi.12121
23. Folmer O., Black M., Hoeh W., Lutz R., Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3: 294-299.
24. GBIF.org. 2025. GBIF (11 June 2025) Occurrence Download https://doi.org/10.15468/dl.venuy5.
25. Goff M.L., 1986. Spider mites (Acari: Tetranychidae) in the Hawaiian Islands. International Journal of Acarology, 12: 43-49. https://doi.org/10.1080/01647958608683437
26. Hoy M.A. 2011. Agricultural Acarology. Introduction to Integrated Mite Management. CRC Press, 410 pp.
27. Hulme P.E. 2009. Trade, transport and trouble: managing invasive species pathways in an era of globalization. Journal of Applied Ecology, 46: 10-18. https://doi.org/10.1111/j.1365-2664.2008.01600.x
28. Kumar S., Stecher G., Suleski M., Sanderford M., Sharma S., Tamura K. 2024. MEGA12: Molecular Evolutionary Genetic Analysis Version 12 for Adaptive and Green Computing, Molecular Biology and Evolution, 41(12), msae263. https://doi.org/10.1093/molbev/msae263
29. Liebhold A.M., Brockerhoff E.G., Garrett L.J., Parke J.L., Britton K.O. 2012. Live plant imports: the major pathway for forest insect and pathogen invasions of the US. Frontiers in Ecology and the Environment, 10: 135-143. https://doi.org/10.1890/110198
30. López-Olmos S., Ferragut F. 2023 The newcomer takes it all: the invader Texas citrus mite, Eutetranychus banksi (Acari: Tetranychidae), displaces the resident relatives in citrus agrosystems. Biological Invasions, 25(10): 3171-3192 https://doi.org/10.1007/s10530-023-03099-z
31. Ma E.P., Yuan Y.L., 1981. New species and new records of tetranychid mites from China II. (Acarina: Tetranychidae). Zoological Research, 2: 281-287.
32. Matsuda T, Kozaki T, Ishii K, Gotoh T 2018. Phylogeny of the spider mite sub-family Tetranychinae (Acari: Tetranychidae) inferred from RNA-Seq data. Plos One 13(9): e0203136. https://doi.org/10.1371/journal.pone.0203136
33. McGregor E.A. 1914. The red spider on citrus. USDA Bureau of Entomology Bulletin, 90: 1-64.
34. McGregor E.A. 1950. Mites of the family Tetranychidae. American Midland Naturalist, 44: 257-420. https://doi.org/10.2307/2421963
35. Mendonca R.S., Navia D., Diniz I.R., Auger P., Navajas M. 2011. A critical review on some closely related species of Tetranychus sensu stricto (Acari: Tetranychidae) in the public DNA sequences databases. Experimental and Applied Acarology, 55: 1-23. https://doi.org/10.1007/s10493-011-9453-5
36. Migeon A., Dorkeld F. 2025. Spider Mites Web: a comprehensive database for the Tetranychidae. Montpellier: INRAE/CBGP; [25 september 2025]. Available from: https://www1.montpellier.inrae.fr/CBGP/spmweb/public/
37. Migeon A., Auger P., Fossati-Gaschignard O., Hufbauer R.A., Miranda M., Zriki G., Navajas M. 2023. Climate of origin influences how a herbivorous mite responds to drought-stressed host plants. Peer Community Journal, 3: e44. https://doi.org/10.24072/pcjournal.272
38. Mushtaq H.M.S., Alatawi FJ, Kamran M., Flechtmann C.H.W. 2021. The genus Oligonychus Berlese (Acari, Prostigmata, Tetranychidae): taxonomic assessment and a key to subgenera, species groups, and subgroups. ZooKeys 1079: 89-127. https://doi.org/10.3897/zookeys.1079.75175
39. Navajas M., Migeon A., Estrada-Peña A., Mailleux A.-C., Servigne P., Petanović R. 2010. Mites and ticks (Acari). Chapter 7.4. In: Roques A. et al. (Eds). Arthropod invasions in Europe. BioRisk 4: 149-192. https://doi.org/10.3897/biorisk.4.58
40. Navajas M., Moraes G.J. de, Auger P., Migeon A., 2013. Review of the invasion of Tetranychus evansi: biology, colonization pathways, potential expansion and prospects for biological control. Experimental and Applied Acarology, 59(1-2): 43-65. https://doi.org/10.1007/s10493-012-9590-5
41. Navia D., Flechtmann C.H.W., Moraes G.J. de, Lindquist E.E. 2011. Review of phytophagous mites (Acari: Prostigmata) from South America. Experimental and Applied Acarology, 54: 57-93. https://doi.org/10.1007/s10493-011-9441-2
42. Ochoa, R., Aguilar, H., Sanabria, C. 1990. Acaros fitoparasitos asociados al cultivo del mango (Mangifera indica L.) en Costa Rica. Manejo Integrado de Plagas, 16: 32-37.
43. Ovalle T.M., Vásquez-Ordóñez A.A., Jimenez J., Parsa S., Cuellar W.J., Lopez-Lavalle L.A.B. 2020. A simple PCR-based method for the rapid and accurate identification of spider mites (Tetranychidae) on cassava. Scientific Reports, 10: 19496. https://doi.org/10.1038/s41598-020-75743-w
44. Pritchard A.E., Baker E.W. 1955. A revision of the spider mite family Tetranychidae. Memoirs https://doi.org/10.5962/bhl.title.150852
45. Series, San Francisco, Pacific Coast Entomological Society, 472 pp.
46. Razuvaeva A.V., Ulyanova E.G., Skolotneva E.S., Andreeva I.V. 2023. Species identification of spider mites (Tetranychidae: Tetranychinae): a review of methods. Vavilovskii Zhurnal Genet Selektsii. 27:240-249. https://doi.org/10.18699/VJGB-23-30
47. Rioja T., Tello V., Zarzar M., Cardemil A., Ceballos, R. 2019. Avocado `Hass' leaf age affects life table parameters of Oligonychus yothersi (McGregor) (Acari: Tetranychidae) under laboratory conditions. Chilean Journal of Agricultural Research, 79: 557-564. https://doi.org/10.4067/S0718-58392019000400557
48. Rode P.A., Alves L.F.A., Loeblein J.S., Toldi M., Ferla N. J. 2023. Systemic activity of azadirachtin on Oligonychus yothersi (Acari: Tetranychidae) on yerba mate plants. Arquivos do Instituto Biológico, 90, e00162022. https://doi.org/10.1590/1808-1657000162022
49. Sheikholeslam-Zadeh S., Sadeghi-Nameghi H. 2010. First records of four mite species (Acari: Tetranychidae) in Iran. Applied Entomology and Phytopathology, 78: 121-126.
50. Torres E., Álvarez-Acosta C., del Pino M., Wong M.E., Boyero J.R., Hernández-Suárez E., Vela J.M. 2023. Economic impact of the Persea mite in Spanish avocado crops. Agronomy, 13(3): 668 https://doi.org/10.3390/agronomy13030668
51. Torres, E., Álvarez-Acosta, C., Ferragut, F., Hernández-Suárez, E.M., 2024. Oligonychus perseae (Tetranychidae) invasion in the Canary Islands: history, management and current situation. Agronomy, 14(5): 920. https://doi.org/10.3390/agronomy14050920
52. Vargas-Madriz A.E., Sánchez-Velázquez O.A., López-Olguín J.F., Sánchez-Sánchez O. 2025. Population fluctuation of phytophagous mites and their impact on blackberry (Rubus spp.) in Jalisco, Mexico. Agronomy, 15: 1970. https://doi.org/10.3390/agronomy15081970
53. Venette R.C., Hutchison W.D. 2021. Invasive Insect Species: Global Challenges, Strategies and Opportunities. Front. Insect Sci. 1: 650520. https://doi.org/10.3389/finsc.2021.650520

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