Skip to content


  • Short report
  • Open Access

Elevation of Pseudoskusea, Rusticoidus and Protomacleaya to valid subgenera in the mosquito genus Aedes based on taxon naming criteria recently applied to other members of the Tribe Aedini (Diptera: Culicidae)

Parasites & Vectors20158:668

  • Received: 8 October 2015
  • Accepted: 8 December 2015
  • Published:



Pseudoskusea, Rusticoidus and Protomacleaya were well-recognized, morphologically distinct subgenera within the genus Aedes prior to a series of taxonomic changes over the past 15 years by Reinert, Harbach and Kitching, when they were recognized as subgenera of the genus Ochlerotatus. In our recent effort to stabilize the Tribe Aedini, we synonymized these subgenera and associated species back into the genus Aedes, but incorrectly assigned them as putative informal groups, instead of reinstating them to subgenera.


Here we formally elevate three traditionally recognized subgenera (Pseudoskusea, Rusticoidus and Protomacleaya) within the genus Aedes.


  • Tribe Aedini
  • Aedes
  • Subgenera
  • Elevation


The tribe Aedini is comprised of about a third of all recognized mosquito species, and includes many vectors of debilitating viral diseases to humans, such as Dengue and Chikungunya. Within the tribe, the genus Aedes, in the traditional sense, is the largest genus in the tribe with 932 species. Other aedine genera are Armigeres, Eretmapodites, Haemagogus, Heizmannia, Opifex, Psorophora, Udaya and Verrallina. During the past 11 years, based on a series of morphological phylogenetic studies by Reinert, Harbach & Kitching (RH&K) [14], and the taxonomic actions resulting from those studies, the original genus Aedes was split into 74 genera, reducing the genus Aedes from over 900 species [58], to only 12. Chief among the reasons given by RH&K to elevate so many genera was the author’s claim of an unreferenced “principle of equivalent rank.” This implied that if traditionally accepted genera were phylogenetically co-equal with other clusters of species in their analyses, the newly recognized groups should also be given similar taxonomic status. These taxonomic actions were highly controversial [9, 10] and resulted in wide-spread confusion about which names to apply to most vectors of disease organisms in genus Aedes (see Table one in [12]). For example, during this period, Aedes japonicus (Theobald), an invasive species and proven vector of West Nile virus and Cache Valley virus, was known variously as Aedes (Finlaya) japonicus [5], Ochlerotatus (Finlaya) japonicus [11], ‘Ochlerotatus’ (‘Finlaya’) japonicus [1] and Hulecoeteomyia japonica [2].

Close scrutiny of the RH&K phylogenetic results and a reanalysis of their dataset led Wilkerson et al. [12] to the conclusion that based on the evidence provided by RH&K the classification changes they promoted and that resulted in the split of the well-known genus Aedes into so many genera, were not warranted. Aedes was therefore reinstated [12], but to preserve their phylogenetic hypotheses the RH&K genera were reduced in rank to subgenera of Aedes. Any subgenera in the fragmented RH&K system were reduced to putative informal group status [12]. Rationalization for reinstatement of genus Aedes to include all “traditionally” accepted species was based on opinions promoting a conservative approach to classification change based on new phylogenetic analyses [1315]. These opinions were comprehensively solidified by Vences et al. [16] who, in detail, discussed the relationship between nomenclatorial utility and phylogenetic accuracy. As a guide to determine the suitability for classification changes they proposed a number of Taxon Naming Criteria (TNCs). Appropriate TNCs were cited to reinstate the “traditional” species in genus Aedes [12]. Central to these arguments reinstating genus Aedes, while retaining other traditional aedine genera were: TNC 2, Clade Stability; TNC 3, Phenotypic Diagnosibility; TNC 8, Manageability; TNC 10, Nomenclatural Stability, and; TNC 11, Community Consensus. Since, to these authors [12], there was no compelling evidence warranting changing the classification of traditional diagnosable genera, the traditional genera in tribe Aedini should be retained until strong, multiple lines of evidence are produced showing the contrary.

Following our recent publication reinstating the genus Aedes [12], we revisited the above rationale and realized that three traditionally recognized Aedes subgenera (Pseudoskusea, Rusticoidus and Protomacleaya), recognized as subgenera by RH&K in their genus Ochlerotatus, were incorrectly synonymized as putative informal groups [12], when they should have been reinstated as bona fide subgenera of the genus Aedes. All are diagnosable, well-known traditional groupings and should be retained as such. Taxonomic information for each subgenus, including important references and component species are given in Appendix.

Conclusion and formal taxonomic action

Here, we formally retrieve Pseudoskusea, Rusticoidus and Protomacleaya from synonymy within the Aedes subgenus Ochlerotatus [12], and elevate all three as subgenera of the genus Aedes.



We thank Bruce Harrison for careful review of the manuscript and his many helpful comments. This research was performed under a Memorandum of Understanding between the Walter Reed Army Institute of Research and the Smithsonian Institution, with institutional support provided by both organizations. This manuscript was prepared whilst YML held a National Research Council (NRC) Research Associateship Award at the Walter Reed Army Institute of Research. The material to be published reflects the views of the authors and should not be construed to represent those of the U.S. Department of the Army or the U.S. Department of Defense.

Open AccessThis article is distributed under the terms of the Creative Commons Attribution 4.0 International License (, which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver ( applies to the data made available in this article, unless otherwise stated.

Authors’ Affiliations

Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA
Walter Reed Biosystematics Unit, Museum Support Center, Smithsonian Institution, Suitland, Maryland, USA
Department of Entomology, Walter Reed Army Institute of Research, Silver Spring, Maryland, USA
Faculty of Preventative Medicine and Biometrics, Uniformed Services University of the Health Sciences, Bethesda, Maryland, USA


  1. Reinert JF, Harbach RE, Kitching IJ. Phylogeny and classification of Aedini (Diptera: Culicidae), based on morphological characters of all life stages. Zool J Linn Soc. 2004;142:289–368. doi:10.1111/j.1096-3642.2004.00144.x.View ArticleGoogle Scholar
  2. Reinert JF, Harbach RE, Kitching IJ. Phylogeny and classification of Finlaya and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages. Zool J Linn Soc. 2006;148:1–101. doi:10.1111/j.1096-3642.2006.00254.x.View ArticleGoogle Scholar
  3. Reinert JF, Harbach RE, Kitching IJ. Phylogeny and classification of Ochlerotatus and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages. Zool J Linn Soc. 2008;153:29–114. doi:10.1111/j.1096-3642.2008.00382.x.View ArticleGoogle Scholar
  4. Reinert JF, Harbach RE, Kitching IJ. Phylogeny and classification of tribe Aedini (Diptera: Culicidae). Zool J Linn Soc. 2009;157:700–94. doi:10.1111/j.1096-3642.2009.00570.x.View ArticleGoogle Scholar
  5. Knight KL, Stone A. A catalog of the mosquitoes of the world (Diptera: Culicidae). 2nd ed. College Park, Maryland: Entomological Society of America; 1977. 611 p.Google Scholar
  6. Knight KL. Supplement to a catalog of the mosquitoes of the world (Diptera: Culicidae). College Park, Maryland: Entomological Society of America; 1978. 107 p.Google Scholar
  7. Ward RA. Second supplement to “A catalog of the mosquitoes of the world” (Diptera: Culicidae). Mosq Syst. 1984;16:227–70.Google Scholar
  8. Ward RA. Third supplement to “A catalog of the mosquitoes of the world” (Diptera: Culicidae). Mosq Syst. 1992;24:177–230.Google Scholar
  9. Savage HM, Strickman D. The genus and subgenus categories within Culicidae and placement of Ochlerotatus as a subgenus of Aedes. J Am Mosq Control Assoc. 2004;20:208–14.PubMedGoogle Scholar
  10. Savage HM. Classification of mosquitoes in tribe Aedini (Diptera: Culicidae): Paraphylyphobia, and classification versus cladistic analysis. J Med Entomol. 2005;42:923–7. doi:10.1093/jmedent/42.6.923. PMID: 16465729.PubMedView ArticleGoogle Scholar
  11. Reinert JF. New classification for the composite genus Aedes (Diptera: Culicidae: Aedini), elevation of subgenus Ochlerotatus to generic rank, reclassification of the other subgenera, and notes on certain subgenera and species. J Am Mosq Control Assoc. 2000;16:175–88.PubMedGoogle Scholar
  12. Wilkerson R, Linton Y-M, Fonseca D, Schultz T, Price D, Strickman D. Making mosquito taxonomy useful: A stable classification of tribe Aedini that balances utility with current knowledge of evolutionary relationships. PLoS One. 2015;10, e0133602. doi:10.1371/journal.pone.0133602.PubMedPubMed CentralView ArticleGoogle Scholar
  13. Wiley EO. An annotated Linnaean hierarchy, with comments on natural taxa and competing systems. Syst Zool. 1979;28:308–37. doi:10.1093/sysbio/28.3.308.View ArticleGoogle Scholar
  14. Wiley EO. Phylogenetics. The Theory of Phylogenetic Systematics. New York: Wiley-Interscience; 1981. 439 p.Google Scholar
  15. Wiley EO, Liebermann BS. Phylogenetic Systematics. Theory and Practice of Phylogenetic Systematics. Hoboken, New Jersey: Wiley-Blackwell; 2011.View ArticleGoogle Scholar
  16. Vences M, Guayasamin JM, Miralles A, de la Riva I. To name or not to name: Criteria to promote economy of change in Linnaean classification schemes. Zootaxa. 2013;3636:201–44. doi:10.11646/zootaxa.3636.2.1.PubMedView ArticleGoogle Scholar
  17. Theobald FV. A monograph of the Culicidae or mosquitoes. London: British Museum (Natural History); 1907. 639 p.Google Scholar
  18. Taylor FH. The Culicidae of Australia. -I. Trans Entomol Soc London. 1914;61:683–708.View ArticleGoogle Scholar
  19. van der Wulp FM. Dipteren aus Neu-Guinea in der Ungarischen National-Museums. Termeszetrajzi Fuzetek, kiadja a Magyar nemzeti Muzeum, Budapest. 1898;21:409–26.Google Scholar
  20. Dobrotworsky NV. Notes on Australian mosquitoes (Diptera, Culicidae). V. Subgenus Pseudoskusea in Victoria. Proc Linn Soc New South Wales; 1961. 257–267.Google Scholar
  21. Dobrotworsky NV. The mosquitoes of Victoria (Diptera, Culicidae). London and New York: Melbourne University Press; 1965. 237 p.Google Scholar
  22. Lee DJ, Dyce AL, O’Gower AK. The Culicidae of the Australasian region, vol. 3. Canberra, Australia: Australian Government Publishing Service; 1984.Google Scholar
  23. Reinert JF. Comparative anatomy of the female genitalia of genera and subgenera in tribe Aedini (Diptera: Culicidae). Part XIII. Ochlerotatus Lynch Arribalzaga. Cont Amer Entomol Inst. 2002;33:1–117.Google Scholar
  24. Surcouf J, Gonzalez-Rincones R. Dipteres piqueurs et suceurs de sang actuellement connus de la Republique de Venezuela. Archives de Parasitologie Paris. 1912;15:248–314.Google Scholar
  25. Schick RX. Mosquito studies (Diptera, Culicidae) XX. The terrens group of Aedes (Finlaya). Cont Amer Entomol Inst. 1970;5:1–158.Google Scholar
  26. Schick RX. Mosquito studies (Diptera, Culicidae) XXIII. Additions and corrections to the revision of the Aedes terrens group. Cont Amer Entomol Inst. 1970;7:13–40.Google Scholar
  27. Zavortink TJ. Mosquito Studies (Diptera, Culicidae) XXVIII. The new world species formerly placed in Aedes (Finlaya). Cont Amer Entomol Inst. 1972;8:1–206.Google Scholar
  28. Shevchenko AK, Prudkina NS. On morphology of genitals in mosquito males from the Aedes genus. Vestnik Zoologii. 1973;6:40–7.Google Scholar
  29. Reinert JF. The subgenus Rusticoidus of genus Aedes (Diptera: Culicidae) in Europe and Asia. Europ Mosq Bull. 1999;4:1–7.Google Scholar


© Wilkerson and Linton. 2015


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate. Please note that comments may be removed without notice if they are flagged by another user or do not comply with our community guidelines.