Eur. J. Entomol. 123: 90-121, 2026 | DOI: 10.14411/eje.2026.011

Integrative taxonomy reveals a hidden diversity: Redescriptions and new records of Rhizoecidae (Sternorrhyncha: Coccomorpha) in GermanyOriginal article

Alejandro CABALLERO ORCID...1, Mehmet Bora KAYDAN ORCID...2, Andrea RAMOS-PORTILLA ORCID...3, Michael J. RAUPACH ORCID...4
1 Faculty of Biology, Ludwig Maximilian Universität, München, Germany and Section Hemiptera, Bavarian State Collection of Zoology (SNSB-ZSM), München, Germany; e-mail: alejandrocab66@gmail.com
2 Biotechnology Development and Research Centre, Çukurova University, Adana, Turkey; e-mail: bkaydan@gmail.com
3 Instituto Colombiano Agropecuario, Bogotá, Colombia; e-mail: andrea.ramos@ica.gov.co
4 Section Hemiptera, Bavarian State Collection of Zoology (SNSB-ZSM), München, Germany; e-mail: raupach@snsb.de

The scale insect family Rhizoecidae in Germany has historically been documented as comprising six species from two genera: Rhizoecus albidus Goux, 1942, Rhizoecus cacticans (Hambleton, 1946), Rhizoecus franconiae Schmutterer, 1956, Ripersiella caesii (Schmutterer, 1956), Ripersiella halophila (Hardy, 1868), and Ripersiella hibisci (Kawai & Takagi, 1971). To assess changes in this biodiversity, we collected 37 samples from potted plants across the botanical gardens of Berlin, Bonn, Frankfurt, Hamburg, and Munich. Species identifications were conducted using integrative taxonomy, combining morphological analysis of slide-mounted specimens, molecular data from the COI region, and ecological information. Our study identified eight species: Geococcus coffeae Green, 1933, Rhizoecus arabicus Hambleton, 1976, Rh. cacticans, Rhizoecus dianthi Green, 1926, Rhizoecus nemoralis (Hambleton, 1946), Rhizoecus falcifer Kunckel d'Herculais, 1878, Rhizoecus simplex (Hambleton, 1946) and Ripersiella aloes (Williams & Pellizzari, 1997). Except for Rh. cacticans, all are newly recorded for Germany, increasing the total German Rhizoecidae fauna to 13 species across three genera. We provide detailed morphometric redescriptions for Rh. arabicus, Rh. franconiae, Rh. simplex, and Ri. caesii based on type material, along with morphological notes for the other species. The study also resolves nomenclatural issues for Rh. franconiae and Ri. hibisci designating Lectotypes and reports 61 new host records for eight species. We present a taxonomic key for the updated list of Rhizoecidae species recorded in Germany. This research significantly revises the understanding of rhizoecid diversity in the country, revealing that the fauna is more than twice as rich as previously known.

Keywords: Root mealybugs, invasive species, barcoding

Received: November 28, 2025; Revised: March 9, 2026; Accepted: March 9, 2026; Published online: March 25, 2026  Show citation

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CABALLERO, A., KAYDAN, M.B., RAMOS-PORTILLA, A., & RAUPACH, M.J. (2026). Integrative taxonomy reveals a hidden diversity: Redescriptions and new records of Rhizoecidae (Sternorrhyncha: Coccomorpha) in Germany. EJE123, Article 90-121. https://doi.org/10.14411/eje.2026.011
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References

  1. Ben-Dov Y. 1994: A systematic Catalogue of the Mealybugs of the World (Insecta, Homoptera, Coccoidea, Pseudococcidae and Putoidae): With Data on Geographical Distribution, Host Plants, Biology, and Economic Importance. Intercept, Andover, UK, 686 pp.
  2. Cox J.M. 1978: Revision of the Rhizoecus species (Homoptera: Pseudococcidae) known from New Zealand. - N. Z. J. Zool. 5: 623-638. Go to original source...
  3. Danzig E.M., Gavrilov I.A. & Trapeznikova I.V. 2008: A new pest from a greenhouse of St. Petersburg, Rhizoecus dianthi Green (Homoptera, Pseudococcidae), with karyotype data. - Entomol. Rev. 88: 676-678. Go to original source...
  4. Danzig E.M. & Gavrilov I.A. 2009: Mealybugs of the genus Rhizoecus Kunckel d Herculais, 1878 (Homoptera: Pseudococcidae) of the fauna of Russia and adjacent countries. - Zoosystematica Ross. 18: 224-245. Go to original source...
  5. Dormann C.F., Gruber B. & Fründ J. 2008: Introducing the bipartite Package: Analysing ecological networks. - R News 8: 8-11.
  6. Felsenstein J. 1985: Confidence limits on phylogenies: An approach using the bootstrap. - Evolution 39: 783-791. Go to original source...
  7. Ferris G.F. 1953: Atlas of the Scale Insects of North America. The Pseudococcidae (Part II). Stanford University Press, Palo Alto, CA, 506 pp.
  8. Fetykó K., Kozár F. & Daróczi K. 2010: Species list of the scale insects (Hemiptera, Coccoidea) of Romania, with new data. - Acta Phytopathol. Entomol. Hung. 45: 291-302. Go to original source...
  9. García Morales M., Denno B., Miller D.R., Miller G.L., Ben-Dov Y. & Hardy N.B. 2016: ScaleNet: A literature-based model of scale insect biology and systematics. - Database 2016: bav118, 5 pp.
  10. Gavrilov-Zimin I.A. 2016: New and earlier unnoted mealybugs and felt scale (Homoptera: Coccinea: Pseudococcidae, Eriococcidae) from Morocco. - Ann. Soc. Entomol. Fr. (NS) 52: 88-94. Go to original source...
  11. Gavrilov-Zimin I.A. & Gapon D.A. 2016: New Afrotropical scale insect pests (Homoptera: Coccinea) under glass in St Petersburg, Russia. - Zoosystematica Ross. 25: 233-238. Go to original source...
  12. Germain J., Matile-Ferrero D., Picart J., Delvare G. & Piron M. 2002: Inventaire des cochenilles présentes sous serre en France et connaissance de leurs parasitoides, un préalable au développement de la lutte biologique. In: Sixth International Conference on Pests in Agriculture, Montpellier, France, 4-6 December 2002. AFPP, Languedoc-Roussillon - Conseil régional, Hérault - Conseil général, ENSAM, INRA, Paris, pp. 253-260.
  13. Giard A. 1897: Sur deux cochenilles nouvelles Ortheziola fodiens nov. spec. et Rhizoecus eloti nov. spec., parasites des racines du caféier à la Guadeloupe. - C. R. Hebd. Séanc. Soc. Biol. 4: 583-585.
  14. Hall T.A. 1999: BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. - Nucleic Acids Symp. Ser. 41: 95-98.
  15. Hambleton E.J. 1946: Studies of hypogeic mealybugs. - Rev. Entomol. Rio Jan. 17: 1-77.
  16. Hambleton E.J. 1973: Florida mealybugs of the genus Rhizoecus with description of a new species (Homoptera: Pseudococcidae). - Proc. Entomol. Soc. Wash. 75: 97-100.
  17. Hambleton E.J. 1976: A revision of the New World mealybugs of the genus Rhizoecus (Homoptera: Pseudococcidae). - U.S.D.A. Tech. Bull. 1522: 1-88.
  18. Hodgson C.J. 2012: Comparison of the morphology of the adult males of the Rhizoecine, Phenacoccine and Pseudococcine mealybugs (Hemiptera: Sternorrhyncha: Coccoidea), with the recognition of the family Rhizoecidae Williams. - Zootaxa 3291: 1-79. Go to original source...
  19. ICZN 1999: International Code of Zoological Nomenclature. The International Trust for Zoological Nomenclature, London, UK, 306 pp.
  20. Jansen M.G.M. & Alferink L.P. 2023: An updated list of scale insects (Hemiptera, Coccomorpha) from import interceptions and greenhouses in the Netherlands. - J. Insect Biodiv. 44: 21-40. Go to original source...
  21. Jansen M. & Westenberg M. 2015: Morphological and molecular studies of a new species of the root mealybug genus Ripersiella Tinsley (Hemiptera: Coccoidea: Rhizoecidae) from greenhouses in the Netherlands and a first incursion of the American root mealybug Rhizoecus keysensis Hambleton in Europe. - Tijdschr. Entomol. 158: 1-19. Go to original source...
  22. Julius Kühn-Institut 2022: Eradication of an Outbreak of Ripersiella hibisci in Germany (Baden-Wuerttemberg). Julius Kühn-Institut, Institute for National and International Plant Health, Braunschweig, 3 pp.
  23. Kimura M. 1980: A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. - J. Mol. Evol. 16: 111-120. Go to original source...
  24. Kosztarab M. & Kozár F. 1988: Scale Insects of Central Europe. Dr. W. Junk and Akadémia Kiado, Budapest, 455 pp. Go to original source...
  25. Kozár F. & Konczné Benedicty Z. 2003: Description of four new species from Australian, Austro-oriental, New Zealand and South Pacific regions (Homoptera, Coccoidea, Pseudococcidae, Rhizoecinae), with a review, and a key to the species Ripersiella. - Boll. Zool. Agrar. Bachic. 35: 225-239.
  26. Kozár F. & Konczné Benedicty Z. 2007: Rhizoecinae of the World. Plant Protection Institute, Hungarian Academy of Science, Budapest, 617 pp.
  27. Kozár F., Szita É., Fetykó K., Neidert D. & Konczné Benedicty Z. 2013: Pajzstetű (Hemiptera: Coccoidea) fajok a magyarországi autópályákon [Species of Hemiptera: Coccoidea on Hungarian Highways]. Institute of Plant Protection of the Hungarian Academy of Sciences, Budapest, 215 pp.
  28. Kozarzhevskaya E. & Reitzel J. 1975: The scale insects (Homo­ptera: Coccoidea) of Denmark. - J. Plant Breed. 79: 1-40.
  29. Kumar S., Stecher, Suleski M., Sanderford M., Sharma S. & Tamura K. 2024: MEGA12: Molecular Evolutionary Genetic Analysis Version 12 for adaptive and green computing. - Mol. Biol. Evol. 41: 1-9. Go to original source...
  30. Künckel D'Herculais J. 1878: Historie de la cochenille vivant sur les racines des palmiers de la section des Seaforthia. Exposé des caractères du genre Rhizoecus. - Ann. Soc. Entomol. Fr. 5: 161-164.
  31. Lindinger L. 1939: Nachtrag zur Schildlaus-fauna Nordwestdeutschlands. - Bombus 10: 37-38.
  32. Malumphy C. & Badmin J.S. 2012: Scale insects and whiteflies (Hemiptera: Coccoidea and Aleyrodoidea) of Watsonian Kent; with a discussion on the impact of naturalised non-native species. - Br. J. Entomol. Nat. Hist. 25: 15-49.
  33. Matile-Ferrero D. & Étienne J. 2006: Cochenilles des Antilles françaises et de quelques autres îles Caraïbes (Hemiptera, Coccoidea) [Cochineal insects of the French Antilles and some other Caribbean islands]. - Rev. Fr. Entomol. 28: 161-190.
  34. Mazzeo G., Longo S., Pellizzari G., Porcelli F., Suma P. & Russo A. 2014: Exotic scale insects (Coccoidea) on ornamental plants in Italy: a never-ending story. - Acta Zool. Bulg. Suppl. 6: 55-61.
  35. McKenzie H.L. 1967: Mealybugs of California with Taxonomy, Biology and Control of North American Species (Homoptera: Coccoidea: Pseudococcidae). University of California Press, Berkeley and Los Angeles, 525 pp. Go to original source...
  36. Moghaddam M. 2013: An annotated checklist of the scale insects of Iran (Hemiptera, Sternorrhyncha, Coccoidea) with new records and distribution data. - ZooKeys 334: 1-92. Go to original source...
  37. POWO 2026: Plants of the World Online. URL: https://powo.science.kew.org/ (last accessed 20 Jan. 2026).
  38. R Development Core Team. 2019: R: A Language and Environment for Statistical Computing. URL: https://www.r-project.org/.
  39. Ramos-Portilla A. & Caballero A. 2022: Rhizoecus spp. (Hemiptera: Coccomorpha: Rhizoecidae). In Kondo T. & Watson G.W. (eds): Encyclopedia of Scale Insect Pests. CAB International, Boston, MA, pp. 121-125.
  40. Ratnasingham S. & Hebert P. 2007: BOLD: The Barcode of Life Data system (www.barcodinglife.org). - Mol. Ecol. Notes 7: 355-364. Go to original source...
  41. Schmutterer H. 1952: Die Ökologie der Cocciden (Homoptera, Coccoïdea) Frankens 1. Abschnitt1. - Z. Angew. Entomol. 33: 369-420. Go to original source...
  42. Schmutterer H. 1956: Neue Rhizoecus-Arten aus Mitteleuropa (Homoptera: Coccoidae: Pseudococcidae). - Beitr. Entomol. 6: 516-521.
  43. Schmutterer H. 1980: Zum Stand der Erforschung der Schildlause (Homoptera, Coccoidea) in der Bundesrepublik Deutschland. [On the status of the scale insect (Homoptera, Coccoidea) in the Federal Republic of Germany.]. - Mitt. Dtsch. Ges. Allg. Angew. Entomol. 2: 49-56.
  44. Simon C., Frati F., Beckenbach A., Crespi B., Liu H. & Flook P. 1994: Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. - Ann. Entomol. Soc. Am. 87: 651-701. Go to original source...
  45. Sirisena U.G., Watson G.W., Hemachandra K.S. & Wijayagunasekara H.N. 2013: A modified technique for the preparation of specimens of Sternorrhyncha for taxonomic studies. - Trop. Agric. Res. 24: 139-149.
  46. Szita É., Blay Goicoechea M.A., París M. & Kaydan M.B. 2022: New data to the scale insect (Hemiptera, Coccomorpha) fauna of Spain. - Acta Phytopathol. Entomol. Hung. 57: 215-219. Go to original source...
  47. Tang F.D. 1992: The Pseudococcidae of China. Shanxi Agricultural University, Taigu, 768 pp.
  48. Williams D.J. 1958: Mealybugs (Pseudococcidae: Homoptera) described by W.M. Maskell, R. Newstead, T.D.A. Cockerell and E.E. Green from the Ethiopian region. - Bull. Br. Mus. Nat. Hist. 6: 205-236.
  49. Williams D.J. 1962: The British Pseudococcidae (Homoptera: Coccoidea). - Bull. Br. Mus. Nat. Hist. 12: 1-79. Go to original source...
  50. Williams D.J. 1985: Australian Mealybugs. British Museum (Natural History), London, 431 pp.
  51. Williams D.J. 2004: Mealybugs of Southern Asia. Natural History Museum, Southdene, Kuala Lumpur, 896 pp.
  52. Williams D.J. & Pellizzari G. 1997: Two species of mealybugs (Homoptera Pseudococcidae) on the roots of Aloaceae in greenhouses in England and Italy. - Boll. Zool. Agrar. Bachic. 29: 157-167.
  53. Williams D.J. & Granara de Willink M.C. 1992: Mealybugs of Central and South America. CAB International, Wallingford, 635 pp.
  54. Zahradník J. 1965: Sur des cochenilles nouvelles dans les serres en Tchécoslovaquie. - Acta Faun. Entomol. Mus. Natl. Pragae 11: 303-306.

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