Allopatric cryptic diversity in the alpine species complex Phtheochroa frigidana s. lat. (Lepidoptera: Tortricidae)

Boyan Zlatkov, Peter Huemer


Allopatric alpine populations of Phtheochroa frigidana s. lat. (Lepidoptera, Tortricidae) are reviewed. In addition to traditional diagnostic characters of external morphology, the genitalia structures of everted vesicae in male genitalia and DNA barcodes are analysed. This new approach supports the existence of five rather than two species in Europe: Phtheochroa schawerdae (Rebel, 1908) comb. nov. (Dinaric Mts, Rila Mts, Pirin Mts) = P. drenowskyi (Rebel, 1916) syn. nov.; P. alpinana sp. nov. (SW Alps); P. apenninana sp. nov. (Apennines); P. frigidana (Guenée, 1845) stat. rev. (Pyrenees) = P. flavidana (Guenée, 1845) = P. sulphurana (Guenée, 1845) = P. andorrana (Millière, 1865); P. cantabriana sp. nov. (Cantabrian Mts). In order to stabilize the nomenclature, a neotype for Eupoecilia frigidana is designated.


Phtheochroa frigidana; species complex; revised taxonomy; DNA barcoding

Full Text:




Anzaldo S.S., Dombroskie J., Brown J.W. 2014. Morphological variation, taxonomic distribution, and phylogenetic significance of cornuti in Tortricinae (Lepidoptera: Tortricidae). Proceedings of the Entomological Society of Washington 116 (1): 1–31.

Bassi G. & Scaramozzino P.L. 1989. Contributo allo studio della microlepidotterofauna Piemontese. 1. Note sul alcuni interessante perperti di Tortricidae (Lepidoptera). Rivista Piemontese di Storia naturale 10: 155–163.

Cracraft J. 1989. Speciation and its ontology. The empirical consequences of alternative species concepts for understanding patterns and processes of differentiation. In: Otte D. & Endler J.A. (eds) Speciation and its Consequences. Sinauer Associates, Sunderland, MA, USA.

de Waard J.R., Ivanova N.V., Hajibabaei M. & Hebert P.D.N. 2008. Assembling DNA barcodes: analytical protocols. In: Cristofre M. (ed.) Methods in Molecular Biology: 275–293. Environmental Genetics. Humana Press Inc., Totowa, NJ, USA.

Gilligan T.M., Baixeras J., Brown J.W. & Tuck K.R. 2014. T@RTS: Online World Catalogue of the Tortricidae (ver. 3.0). Available from [accessed 26 Nov. 2016].

Guenée A. 1845a. Essai sur une nouvelle classification des microlépidoptères et catalogue des espèces européennes connues jusqu’à jour. Annales de la Société entomologique de France 2 (3): 297–344. Available from [accessed 16 Oct. 2017].

Guenée A. 1845b. Europaeorum Microlepidopterorum Index Methodicus, sive Pyrales, Tortrices, Tineae et Alucitae Linnaei, secundum novum naturalemque Ordinem dispositae, Nominibus Genuinis restitutis, Synomimia accurate elucidata, Locis indicatis, novisque Speciebus aut Larvis brevi descriptis. Pars prima: Sistens Tortrices, Phycidas, Crambidas, Tinearumque initiumaut Larvis brevi descriptis. Pars prima: Sistens Tortrices, Phycidas, Crambidas, Tinearumque initium. Roret, Paris, France.

Hebert P.D.N., Cywinska A., Ball S.L. & de Waard J.R. 2003. Biological identifications through DNA barcodes. Proceedings of the Royal Society B 270: 313–321.

Hebert P.D.N., de Waard J.R. & Landry J.-F. 2009. DNA barcodes for 1/1000 of the animal kingdom. Biology Letters 6: 359–362.

Huemer P. 2015. DNA-Barcoding von Schmetterlingen – alpine Artenvielfalt neu bewertet. Acta ZooBot Austria 152: 161–165.

Huemer P. & Hebert P.D.N. 2011. Cryptic diversity and morphology of high alpine Sattleria – a case study combining DNA barcodes and morphology (Lepidoptera: Gelechiidae). Zootaxa 2981: 1–22.

Huemer P. & Mutanen M. 2015. Alpha taxonomy of the genus Kessleria Nowicki, 1864, revisited in light of DNA-barcoding (Lepidoptera, Yponomeutidae). ZooKeys 503: 89–133.

Kandul N.P., Lukhtanov V.A., Dantchenko A.V., Coleman J.W.S., Sekercioglu C.H., Haig D. & Pierce N.E. 2004. Phylogeny of Agrodiaetus Hübner, 1822 (Lepidoptera: Lycaenidae) inferred from mtDNA sequences of COI and COII and nuclear sequences of EF1-α: karyotype diversification and species radiation. Systematic Biology 53 (2): 278–298.

Kovács Z. & Kovács S. 2004. The occurrence of Phtheochroa drenowskyi (Rebel, 1916) (Lepidoptera, Tortricidae, Cochylini) in Romania. Entomologica Romanica 8–9: 39–42.

Lesar T., Habeler H. & Arenberger E. 2009. Prispevek k poznavanju metuljev (Lepidoptera) Slovenije II: nove vrste metuljčkov (Microlepidoptera). Natura Sloveniae 11 (2): 39–60.

Millière P. 1845. Iconographie et Description de Chenilles et Lépidoptères inédits 2. F. Savy, Paris.

Mutanen M., Hausmann A., Hebert P.D.N., Landry J.-F., de Waard J.R. & Huemer P. 2012. Allopatry as a Gordian knot for taxonomists: patterns of DNA barcode divergence in arctic-alpine Lepidoptera. PLoS One 7 (10): e47214.

Pazhenkova E.A., Zakharov E.V. & Lukhtanov V.A. 2015. DNA barcoding reveals twelve lineages with properties of phylogenetic and biological species within Melitaea didyma sensu lato (Lepidoptera, Nymphalidae). ZooKeys 538: 35–46.

Ragonot E.-L. 1894. Notes synonymiques sur les microlépidoptères et description d’espèces peu connues ou inédites. Annales de la Société entomologique de France 63: 161–226. Available from [accessed 16 Oct. 2017].

Ratnasingham S. & Hebert P.D.N. 2007. BOLD: The Barcode of Life Data System ( Molecular Ecology Notes 7: 355–364.

Ratnasingham S. & Hebert P.D.N. 2013. A DNA-based registry for all animal species: the Barcode Index Number (BIN) system. PLoS One 8: e66213.

Razowski J. 1970. Cochylidae. In: Amsel H.G., Gregor F. & Reisser H. (eds) Microlepidoptera Palaearctica 3. Georg Fromme, Vienna, Austria.

Razowski J. 1991a. The catalogue of the species of Tortricidae (Lepidoptera). Part I: Palaearctic Chlidanothinae and Tortricinae: Cochylini, Tortricini, Ceracini, Cnephasiini. Acta Zoologica Cracoviensia 34 (1): 99–162.

Razowski 1991b. Notes on Cochylinae genus Phtheochroa Stephens (Lepidoptera: Tortricidae) with descriptions of new American species. Acta Zoologica Cracoviensia 34 (1): 163–187.

Razowski J. 2002. Tortricidae of Europe. Volume 1. Tortricinae and Chlidanothinae. František Slamka, Bratislava, Slovakia.

Razowski J. 2008. Tortricidae of the Palaearctic Region. Volume 1. Tortricini and General Part. František Slamka, Bratislava, Slovakia.

Razowski J. 2009. Tortricidae of the Palaearctic Region. Volume 2. Cochylini. František Slamka, Bratislava, Slovakia.

Rebel H. 1916. Beitrag zur Lepidopterenfauna Bulgariens. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien 66: 36–42.

Robinson G. 1976. The preparation of slides of Lepidoptera genitalia with special reference to the Microlepidoptera. Entomologist’s Gazette 27: 127–132.

Schawerda K. 1908. Bericht der Sektion für Lepidopterologie. Verhandlungen der Kaiserlich- Königlichen Zoologisch-Botanischen Gesellschaft in Wien 58: 250–273.

Tamura K., Stecher G., Peterson D., Filipski A. & Kumar S. 2013. MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution 30: 2725–2729.

Trematerra P. 2003. Catalogue of Lepidoptera Tortricidae of the Italian fauna: geonomy, Italian distribution, biological notes, identification. Bollettino di Zoologia Agraria e di Bachicoltura, Series II, 35 (Suppl.): 1–270.

Varga Z.S. & Schmitt T. 2008. Types of oreal and oreotundral disjunctions in the western Palearctic. Biological Journal of the Linnean Society 93: 415–430.

Zlatkov B. 2011. A preliminary study of everted vesicae of several leafrollers (Tortricidae). Nota Lepidopterologica 33: 285–300. Available from [accessed 16 Oct. 2017].

Zlatkov B. & Huemer P. 2016. Phtheochroa unionana (Kennel, 1900) recognised as a dimorphic Cochylini species, with description of the hitherto unknown male genitalia (Lepidoptera, Tortricidae). Nota Lepidopterologica 39 (2): 113–121.


  • There are currently no refbacks.

License URL: