Taxonomic comments on the treatment of the Zygaenidae (Lepidoptera) in volume 3 of Moths of Europe, Zygaenids, Pyralids 1 and Brachodids (2012)

Critical taxonomic comments are provided on the section dealing with the Zygaenidae in volume 3 of Moths of Europe (Leraut 2012). A number of newly described nominal taxa in that work are here synonymised as are nominal subgenera and subspecies that were reinstated as valid. At the subgeneric level these are Hesychia Hübner, [1819] (syn. rev.), Coelestis Burgeff, 1926 (syn. rev.) and Santolinophaga Burgeff, 1926 (syn. rev.), all synonyms of the monophyletic subgenus Mesembrynus Hübner, [1819], and Coelestina Holik, 1953 (syn. rev.), Epizygaena Jordan, 1907 (syn. rev.) and Lictoria Burgeff, 1926 (syn. rev.), all synonyms of the paraphyletic subgenus Agrumenia Hübner, [1819]. At the subspecific level Adscita geryon parisiensis Leraut, 2012 (syn. n.) and A. geryon aeris (Verity, 1946) (syn. rev.) are synonyms of A. geryon geryon (Hübner, [1813]). Zygaena exulans altaretensis Le Charles, 1942 (syn. rev.) and Z. exulans bourgognei Le Charles, 1942 (syn. rev.) are synonyms of Z. exulans exulans (Hohenwarth, 1792). Zygaena trifolii vindilisensis Leraut, 2012 (syn. n.) is a synonym of Z. trifolii subsyracusia Verity, 1925. Zygaena carniolica besseensis Leraut, 2012 (syn. n.) and Z. carniolica rogervillensis Leraut, 2012 (syn. n.) are synonyms of Z. carniolica modesta Burgeff, 1914. Zygaena hilaris nigriventris Leraut, 2012 (syn. n.) is a synonym of Z. hilaris chrysophaea Le Charles, [1934]. Zygaena rhadamanthus boixolsis Aistleitner, 1990 (syn. n.) is a synonym of Z. rhadamanthus rhadamanthus Esper, [1789], Z. rhadamanthus cleui Dujardin, 1956 (syn. rev.) is a synonym of Z. rhadamanthus grisea Oberthür, 1909, and Z. rhadamanthus aragonia Tremewan, 1961 (stat. rev.), Z. rhadamanthus azurea Burgeff, 1914 (stat. rev.) and Z. rhadamanthus aurargentea Mazel, 1979 (stat. rev.) are reinstated as valid subspecies. Résumé. Les auteurs font une analyse critique de la taxonomie employée pour le chapitre traitant des Zygaenidae, dans le volume 3 de la série «papillons de nuit d’Europe». Un certain nombre de taxons nominaux nouvellement décrits sont mis en synonymie, ainsi que des sous-genres et sous-espèces qui avaient été considérés comme valides. Au niveau subgénérique c’est le cas pour Hesychia Hübner, [1819] (syn. rev.), Coelestis Burgeff, 1926 (syn. rev.) et Santolinophaga Burgeff, 1926 (syn. rev.), qui doivent être replacés dans le sous-genre monophylétique Mesembrynus Hübner, [1819] et Coelestina Holik, 1953 (syn. rev.), Epizygaena Jordan, 1907 (syn. rev.) et Lictoria Burgeff, 1926 (syn. rev.), qui doivent eux être placés dans le sous-genre paraphylétique Agrumenia Hübner, [1819]. Pour ce qui concerne les sous-esNota Lepi. 37(2) 2014: 123–133 | DOI 10.3897/nl.37.7940 Efetov et al.: Taxonomic comments on the treatment of the Zygaenidae... 124 pèces, il s’agit de Adscita geryon parisiensis Leraut, 2012 (syn. n.) et A. geryon aeris (Verity, 1946) (syn. n.), qui doivent être rattachés à A. geryon geryon (Hübner, [1813]); Z. exulans altaretensis Le Charles, 1942 (syn. rev.) et Z. exulans bourgognei Le Charles, 1942 (syn. rev.) synonymes de Z. exulans exulans (Hohenwarth, 1792); Z. trifolii vindilisensis Leraut, 2012 (syn. n.) synonyme de Z. trifolii subsyracusia Verity, 1925; Z. carniolica besseensis Leraut, 2012 (syn. n.) et Z. carniolica rogervillensis Leraut, 2012 (syn. n.) synonymes de Z. carniolica modesta Burgeff, 1914; Z. hilaris nigriventris Leraut, 2012 (syn. n.) synonyme de Z. hilaris chrysophaea Le Charles, [1934]; enfin Z. rhadamanthus boixolsis Aistleitner, 1990 (syn. n.) synonyme de Z. rhadamanthus rhadamanthus (Esper, [1789]), Z. rhadamanthus cleui Dujardin, 1956 (syn. rev.) synonyme de Z. rhadamanthus grisea Oberthür, 1909. Zygaena rhadamanthus aragonia Tremewan, 1961 (stat. rev.), Z. rhadamanthus azurea Burgeff, 1914 (stat. rev.) et Z. rhadamanthus aurargentea Mazel, 1979 (stat. rev.), sont réintégrées comme sous-espèces valides.


Introduction
In a recent review of the Zygaenidae of Europe (Leraut 2012), a number of questionable taxonomic changes were proposed, many of which lack convincing morphological and/or phylogenetic support (Efetov et al. 2013).For example, a number of subgenera within the genus Zygaena were unjustifiably reinstated as valid, as were a number of taxa at subspecific level.Moreover, five subspecies and several infrasubspecific forms were newly described, but only the former are dealt with in the present paper, as infrasubspecific forms have no status under the International Code of Zoological Nomenclature (ICZN 1999).In order to obtain a clear understanding of the problem, it is recommended that the review (Efetov et al. 2013) of the handbook and the present article are read together.

Phaudidae
Although treated as a subfamily of the Zygaenidae by Leraut (2012: 44), the Phaudidae were ele- vated to full family status within the Zygaenoidea by Niehuis et al. (2006: 822, fig.3), a placement that is now widely accepted by lepidopterists (van Nieukerken et al. 2011) and fully supported by the present authors.
Adscita geiyon razza aeris Verity, 1946, Redia 31: 154.Type-locality.France: [Alpes-Maritimes], Saint-Bamabé.Syn.rev.Adscita geiyon parisiensis Leraut, 2012 Distribution and taxonomic notes.Adscita geryon geryon is distributed from the Iberian Peninsula and Britain to European Russia, the Crimea and Turkey.Leraut (2012: 61 ) reinstated the nominal taxon A. geryon aeris (Verity, 1946) as valid and on the following page newly described a subspe- cies from the Paris region.It is acknowledged that^.geryon is an extremely variable species, both in phenotype and genitalic morphology, but we see no justification for recalling a subspecies from synonymy, where it was placed by Efetov andTarmann (1999: 28, 2012: 3 1), or in describing a new subspecies from France.Accordingly, both taxa are here formally placed as synonyms (syn.rev.; syn.n.) of the nominotypical subspecies A. geryon geryon.

Zygaeninae
Subgenera.In discussing the classification of the genus Zygaena Fabricius, 1775, at the subgeneric level, Leraut (2012: 67-68) refers to a paper by Niehuis et al. (2006) in which the subge- nera are not mentioned.Presumably the intention was to refer to the evolutionary history of the genus, as based on nuclear and mitochondrial DNA-sequencing by Niehuis et al. (2007).The latter paper provides a phytogeny in which the Zygaena species are placed into species-groups within three subgenera, viz.Mesembrymis Hübner, [1819], Agrumenia Hübner, [1819], and Zygaena Fabricius, 1775, based on the classification of Alberti (1958Alberti ( , 1959) ) and, supported by their own research, followed by Naumann and Tremewan (1984) and Hofmann andTremewan (1996, 2010).However, Leraut (2012: 67-68) resurrected various nominal subgenera from synonymy and applied them to some of the species groups, viz.Hesychia Hübner, [1819], Coelestis Burgeff, 1926, Santolinophaga Burgeff, 1926, all formerly placed as synonyms of the monophyletic subgenus Mesembrynus Hübner, [1819], and Coelestina Holik, 1935, Epizygaena Jordan, [1907], and Lietoria Burgeff, 1926, all formerly placed as synonyms of Agrumenia Hübner, [1819] (Hofmann and Tremewan 2010).It is accepted that the subgenera Agrumenia and Zygaena are paraphyletic, but Mesembrynus is monophyletic, based on morphology (Al-  berti 1958, 1959), larval host-plants (Hofmann and Tremewan 1996) and DNA analysis (Niehuis et al. 2007).Therefore, to split the last-mentioned subgenus into four subgenera is illogical and artificial, as two {Mesembrynus and Santolinophaga) will then become paraphyletic groupings.As a consequence, the nominal subgeneric taxa Hesyehia, Santolinophaga and Coelestis are here formally reinstated as synonyms (syn.rev.) of Mesembrynus, and Epizygaena, Coelestina and Lietoria are reinstated as synonyms (syn.rev.) of Agrumenia.The full synonymy of the subgenera can be found in Hofmann and Tremewan (1996,2010) and only those relevant to Subspecies.In their systematic catalogue of the Zygaeninae, Hofmann and Tremewan (1996) attempted to provide a classification in which some rationale could be brought to the genus Zygaena Fabricius, 1777, with reference to subspecies; hence an enormous number of subspecific taxa were placed as synonyms.It would appear that the classification in that catalogue has largely been ignored, as Leraut reinstated two subspecific taxa as valid and newly described four within Zygaena.While this would have been acceptable during the first half of the century, the description of new subspecies in the genus Zygaena from mainland Europe at the present time neither reflects current thinking nor the presently accepted concept of a subspecies and is reminiscent of the taxonomy practised in the 1920s and 1930s (e.g.Verity (1925Verity ( , 1926) ) described and named three 'subspecies' of Z. trifolii from England, based on single colonies).Mayr (1969: 41) contended that a subspecies is 'an aggregate of phenotypically similar populations of a species, inhabiting a geographic subdivision of the range of a species, and differing taxonomically from other populations of the species' and that it may consist of many local populations all of which, though very similar, are slightly different from each other genetically and phenotypically.In contrast to the reinstatement of two subspecies, Leraut   (2012) synonymised four without providing justification.All of these nominal subspecific taxa are discussed in detail below.As with the subgenera, the full synonymy of the subspecies can be found in Hofmann and Tremewan (1996) and only those relevant to the field guide (Leraut   2012) are listed below.
©Societas Europaea Lepidopterologica; download unter http://www.biodiversitylibrary.org/ und www.zobodat.atespecially in the females, which is reminiscent of Zygaena rhadamanthus grisea from south-eastern and southern-central France (see above), and by the presence of a strong abdominal cingulum that is also present ventrally.Leraut (2012: 98) placed Zygaena rhadamanthus aurargentea as a synonym of Zygaena rhadamanthus cleui, which he reinstated as a valid subspecies, but we see no justification for this (see also above).The former is here reinstated (stat.rev.) as a valid subspecies, based on its extreme phenotype, which is so strongly different from that of the nominotypical taxon Z rhadamanthus rhadamanthus (Hofmann and Tremewan 1996: 131).
Zygaena rhadamanthus aragonia Tremewan, 1961, The Entomologist's Record and Journal of Variation 73: 4. Type-local- ity.Spain: Teruel, Albarracin. [Holotype S, paratypes 16 (S', 11 $ examined.]Distribution and taxonomic notes.Zygaena rhadamanthus aragonia is distributed in the Spanish provinces of Cuenca and Teruel.While it is reminiscent of Zygaena rhadamanthus grisea from south-eastern and southern-central France, the placement of this taxon as a synonym of Zygaena rhadamanthus cleui by Leraut (2012: 98), which he reinstated as a valid subspecies, is inconsistent with the geographical distribution, its 'griseoid' phenotype and the presence of a strong abdominal cingulum.As a consequence, Zygaena rhadamanthus aragonia is here reinstated (stat.rev.) as a valid subspecies (Hofmann and Tremewan 1996: 130).It should be noted that Zygaena rhadamanthus aurargentea also has a griseoid phenotype and a strong abdominal cingulum, but the two taxa are geographically separated by a large lowland area that includes the River Ebro.
Distribution and taxonomic notes.Zygaena trifolii varies in phenotype from year to year, even within the same colony (A.Hofmann and W. G. Tremewan pers.obs.).Hence, specimens taken from the same island (Belle-Ile-en-Mer) in different years may not match the original description (E.Drouet pers.comm.).On this basis, Zygaena trifolii vindilisensis is synonymised (syn.n.)   with Zygaena trifolii subsyracusia, a subspecies that is distributed along the coastal regions of north-western France, from Loire-Atlantique to Côtes-d'Armor and Ille-et-Villaine, and is also found in the Channel Islands (Hofmann and Tremewan 1996: 185).
General taxonomic comments Leraut (2012: 67) stated that the phylogenetic sequence in the presentation of the species follows Niehuis et al. ( 2007), yet the sister species Z. trifolii and Z. lonicerae are divided by ©Societas Europaea Lepidopterologica; download unter http://www.biodiversitylibrary.org/ und www.zobodat.atZ filipendulae\ the first two species are more closely related to each other than either is to Z. filipendulae and together they form a sister group to the latter (Niehuis et al. 2007; Hofmann  and Tremewan 2010: 123).
The position (Leraut 2012: 68) of Z. persephone in the 'subgenus' Santolinophaga (a syno- nym of Mesembrynus, as discussed above) is also incorrect.While we are aware that the sys- tematic position of Z. persephone is at present enigmatic, we cannot see any justification for placing a species whose larva feeds on Vieia glauea C. Presl.(Fabaceae) (Barragué 1986: 316;Tremewan 1989) in a monophyletic subgenus consisting solely of Apiaceae-, Asteraceae-and Lamiaceae-feeders.No doubt Leraut had difficulty assigning it to a subgenus because it was not dealt with by Niehuis et al. (2007).However, until the taxon has been subjected to DNA analysis, it is better to keep the status quo, i.e. to leave it in the subgenus Zygaena, following Alberti (1958: 280, 313), Naumann and Tremewan (1984: 168) and Hofmann and Tremewan (1996: 142, 2010: 123).
While it is correctly stated (Leraut 2012: 100) that Z. oxytropis is closely similar to Z. rhad- amanthus, there is little evidence that the former could only be a subspecies of the latter and that 'molecular biology' could be used to investigate further.In spite of the fact that a hybrid zone between the taxa occurs (or occurred) in north-western Italy (Burgeff 1951: 11), a phenomenon that is not unusual between the ranges of two closely related Zygaena species, the heterospecificity of Z. oxytropis and Z. rhadamanthus has been confirmed by DNA analysis (Niehuis et al. 2007).
With reference to the treatment of Z. transalpina transalpina and Z. transalpina hippoerepidis (Leraut 2012: 102-106), one has to acknowledge that the relationship between these two taxa has been controversial for many years.Leraut (2012) followed the intensive study by Mazel (2009a, 2009b, 2010) who contends that the genitalic morphology has 'confirmed' that both taxa are valid species and that a hybrid zone between the two taxa occurs in eastern France.However, while Mazel should be acknowledged for the enormous amount of research into the problem, his arguments based on genitalic morphology alone are unconvincing.It should be noted that there are also hybrid populations in Germany where they form a mosaic in their distribution rather than a hybrid zone or tension zone (Hofmann 1994: 285-288).Moreover, DNA analysis by Niehuis et al. (2007) supports their conspecificity, as does the work of Hille (2012) who has recently assessed populations by using genogeographic clustering, based on phenotype, genotype and haplotype variation.New molecular data provided by Hille, based on 200 specimens from Germany, Austria, Slovenia, Croatia, Czech Republic and Slovakia,   augment the sequence data base available for this approach.Further support has been provided by a Bayesian phylogenetic analysis (von Reumont et al. 2012: 45, fig.4).