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Research Article
A synonymic list of names associated with western Palaearctic Melitaea phoebe (Denis & Schiffermüller, 1775) species group taxa (M. phoebe; M. punica Oberthür, 1876; M. ornata Christoph, 1893) (Lepidoptera, Nymphalidae)
expand article infoPeter Russell, W. John Tennent
‡ Natural History Museum, London, United Kingdom
Open Access

Abstract

Following indecision and confusion in the literature regarding nomenclature and distribution of Melitaea phoebe (Denis & Schiffermüller, 1775) and its closely associated congeners M. punica Oberthür, 1876, and M. ornata Christoph, 1893, a synonymic list of phoebe names, and of names both correctly and mistakenly associated with phoebe species-group taxa, is presented. Explanatory footnotes provide the basis of a stable source for future discussion of M. phoebe species-group populations throughout the species’ ranges.

Introduction

According to a recent revision (van Oorschot and Coutsis 2014), the genus Melitaea Fabricius, 1807, comprises some 98 species of phenotypically similar, medium sized nymphaline butterflies that occur throughout most of the Palaearctic Region and beyond. Adults are invariably orange-brown on the upper surface, with a series of black lines and other markings; several Melitaea species are notoriously difficult to separate using wing morphology due to their similarity in appearance (Jugovic and Koren 2014).

In this paper, we consider the common and widespread butterflies Melitaea phoebe (Denis & Schiffermüller, 1775) and M. ornata Christoph, 1893; the latter was not fully recognised as a species distinct from M. phoebe until 2005 (see below). We also confirm, not for the first time, that M. punica Oberthür, 1876, historically considered a subspecies of M. phoebe, is restricted in distribution to Africa north of the Sahara; the specificity of M. punica was confirmed by Tóth and Varga (2011). Some fundamental confusion has encompassed the use of names relating to these three phenotypically similar Melitaea species in the western Palaearctic. We believe that confusion, for which the present authors must take some responsibility (with others!), arose as a direct result of the realisation that a second phoebe-like taxon occurred in Europe, before the extent of the range of M. ornata was fully understood. Before that was established, some other names were briefly used in the literature (e.g. emipunica [by Russell et al. 2005] and ogygia [by Varga et al. 2005]).

As recognised here, the western Palaearctic Region extends from the Iberian Peninsula and Africa north of the Sahara in the west to the Ural Mountains and Kazakhstan in the east (approximately 60º East), and from the North Cape of Norway in the north to the Middle East, including Iran and Iraq, in the south (approximately 30º North). We recognise that a number of names relate to Melitaea taxa east of the Urals, and where these apply or may apply to the taxa considered in this paper, they are also included.

Background

A detailed analysis of the Palaearctic forms and varieties associated with Melitaea Fabricius, 1807, was made by Higgins (1941, 1944 [errata], 1955 [additions]). He recognised M. phoebe as a distinct species (Higgins 1941: 325–343, plate 14, figs 1–12; plate 15, fig. 8) with three subspecies: nominotypical phoebe (throughout the region except the Iberian Peninsula and North Africa), occitanica Staudinger, 1871 (Iberian Peninsula) and punica (North Africa). Considering what was available to Higgins at that time, he provided what remains a generally accurate assessment of the division of M. phoebe sensu lato. Higgins provided (1941: 325–343) an annotated list of 74 named forms, mostly originating from the western Palaearctic, and many described individual variations, seasonal forms and aberrations. He later added (Higgins 1955: 118) five additional synonyms for M. phoebe.

So where did it all go so wrong? Modern confusion seems to stem from Hesselbarth et al. (1995: 1030), who mistakenly (see Appendix: Note 3), used the complex name: ‘M. (phoebe) punica telona’ for those phoebe-like butterflies from Turkey which were not, in their opinion – and undoubtedly correctly – M. phoebe sensu stricto. Understandably, their work was used as an anchor for subsequent papers on the region; for example, Çalişkan and Bozaci (2015) described a male aberration of what they considered to be M. phoebe from the province of Adana, Turkey, placing the name ornata as a synonym of “M. punica telona”. They referenced Hesselbarth et al. (1995), from where their use of the combination ‘punica telona’ presumably originated. With several researchers working in different parts of Western Europe and the Russian Federation, it is perhaps understandable that this spurious use of the name punica became so widely used (see Appendix: Note 3).

M. ornata was first recognised as a species separate from M. phoebe simultaneously by Varga et al. (2005), from Hungarian specimens, under the specific name M. ogygia Fruhstorfer, 1908 [TL: Island of Poros, Peleponnese, Greece] and by Russell et al. (2005), from specimens reared from a female from Montagna Longa, Palermo district, Sicily, under the name M. emipunica Verity, 1919 [TL: Palermo district, Sicily, Italy]. The presently known eastern limit of the distribution of the invariably univoltine species M. ornata may coincide with the eastern limit of our interpretation of the western Palaearctic (see above), although a recent publication by Korb et al. (2015) recorded M. ornata from Middle Asia (Kyrghyz Mts., Transili Alatau Mts. and Kungey Ala-Too Mts.). Previously, Korb (2011: 178) reported this same material as M. phoebe saturata but following molecular analysis of the preserved specimens by Korb et al. (2015) their identity was reassessed as M. ornata. We note that larvae from this area are yet to be observed and believe further research is required to confirm their identity.

Placement of M. ornata and some associated Russian taxa as synonyms of M. phoebe by van Oorschot and Coutsis (2014: 60) compounded the confusion referred to above. In the expectation of resolving some long-standing matters of nomenclature and distribution, we present an alphabetical list of names associated with Melitaea phoebe species-group taxa and place each as a synonym of M. phoebe Notes 1 & 6, M. ornata Notes 2 & 6, or M. punica Notes 3 & 6. These actions are explained and supported by detailed notes (see the Appendix). The present authors do not recognise any subspecific divisions of M. ornata or M. punica; however, M. phoebe s.s. occurs in two distinct subspecies based on larval colour and distribution: M. phoebe phoebe Note 4 and M. phoebe occitanica Note 5, with which some names will be synonymised.

Distribution of Melitaea phoebe, M. punica, and M. ornata within the western Palaearctic, as currently known and understood

Distribution of M. phoebe phoebe and M. phoebe occitanica in the western Palaearctic is presented in Note 1. However, it is considered important to clarify some details as to where, so far as is currently known, M. phoebe has been recorded as being present in the literature but is not in fact present:

(1) M. phoebe has been recorded from Lésvos and Chíos (Gaskin and Littler 1986) but it is M. ornata which is present there (Russell and Pateman 2013c); in fact it is almost certain that all reports of M. phoebe from the Aegean Islands should be regarded as M. ornata (Russell and Pamperis 2011, 2012).

(2) In Greece M. phoebe has not been confirmed as occurring south of Mt. Vardousia, Fokida (Lafranchis pers. obs., ex Pamperis pers. comm.), and is absent from the Peleponnese (Lafranchis 2007).

(3) In Italy M. phoebe is not known to occur in southern Calabria, or in Sicily where only M. ornata is present (Russell and Pateman 2011: 26, as telona); however, Villa et al. (2009: 244) gave the distribution of M. phoebe as throughout Italy including Sicily.

(4) In the east M. phoebe is absent from Israel, except coastal areas in the North (Benyamini, pers. comm.), southern and eastern Iran and Iraq, except for one record from the extreme northeast (Tshikolovets et al. 2014: 319), although this may refer to the record of Wiltshire (1957: 33), who referred to f. dorae and stated that there was only one generation per year, flying in early summer. This suggests that this was most probably not M. phoebe but M. ornata.

(5) So far as the authors are aware M. phoebe is also absent from Jordan, where it is replaced by M. ornata (Katbeh-Bader et al. 2003: 17; Wahlberg and Zimmermann 2000: voucher specimen).

(6) It is quite likely that M. phoebe is absent from Syria; although Riemis (1993: 93) recorded M. phoebe from 50 km south of Aleppo on the road to Damascus, this was before M. ornata was separated at species level. The only figured specimens known to the authors from Syria (Van Haeringen 2015) are eight individuals labelled “M. phoebe telona” (= M. ornata), originating from Bloudan (26.iv.2008), Halbourn (27.iv.2008) and Damascus (5.iv.2010). These specimens exhibit antennal and wing morphological characters typical of M. ornata (see Table 1).

Identification difficulties arise in part because of a lack of clear diagnostic features to guarantee separation of adult butterflies; the only apparently constant feature appears to be the colour of late instar larvae. However, there are other features which might aid identification, presented here with an indication of their level of usefulness.

Character M. phoebe M. ornata M. punica Reliability of character
number of ova in batch usually more than 100 usually 30–60 data lacking good
larva L4- final instar head colour black red-brown black confirms M. ornata
final instar larva lateral stripe colour white (phoebe phoebe) orange (phoebe occitanica) no obvious stripe orange good (confirms occitanica outside North Africa)
distal end of antenna club shaped spatulate variable fair
shape of forewing apex acute rounded rounded fair
wing underside background colour creamy white white fair
hindwing underside premarginal marks arcuate triangular variable, often triangular poor
premarginal markings touching veins yes no variable, often not touching fair
voltinism single to triple brooded strictly univotine double to triple brooded good

Those European regions in which both M. phoebe and M. ornata have been recorded as being present (although not necessarily sympatric or synchronic) are as follows: France (Var only), Italy (northern Calabria as far north as Campania (Russell and Pateman 2011), Greece (central and north, see above), Macedonia (FYROM) (Verovnik et al. 2010; Verovnik 2012; Russell et al. 2015), Montenegro (Russell 2015), Slovenia (Russell et al. 2014), Hungary (Varga et al. 2005). There are additional reports of the presence of M. ornata (unconfirmed) from regions where M. phoebe is also known to occur – Croatia (Koren and Štih 2013), Romania (Rákosy and Varga 2001; Székely 2008), Bulgaria (Kolev 2015), and Slovakia (Zitnan pers. comm.). The report by Jakšić (2011: 46–47) of M. ornata from Serbia is considered to be dubious; it is not otherwise known from there, and M. phoebe is widespread throughout that country. In the east, both species occur in Lebanon and Israel (M. phoebe occurs in northern coastal district only; Benyamini, pers. comm.), Turkey (Hesselbarth et al. 1995), the Caucasus (Tshikolovets and Nekrutenko 2012: 293–295; Tikhonov and Russell 2015), the Russian Federation (Russell and Kuznetsov 2012), Syria (but see above), northeast Iraq and northern and western Iran (Tshikolovets et al. 2014). Eisenstein (2000: 190, fig. 234) figured a larva in Israel with a red-brown head feeding on Centaurea iberica (Spreng) (M. ornata: see also Russell et al. 2007).

The authors see no evidence to support subdivision of M. ornata into five subspecies (Tshikolovets 2011: 498–499); previous gaps in the known distribution of this species are rapidly being filled, making recognition of subspecies on a geographical basis increasingly difficult to support. Also, the diverse variety of host-plants used by M. ornata is more likely to be dependent on which Asteraceae species are available for use by larvae in any particular locality, rather than any evolutionary preference resulting in development of subspecies. The differing ability of closely related species to metabolise apparently suitable host-plants is also significant (Tóth et al. 2015); however, it is of interest to note that different host-plants were being used by M. ornata on the adjacent eastern Aegean Islands of Lésvos and Chíos, where adult butterflies were almost identical (Russell and Pamperis 2011; Russell and Pateman 2013c).

Despite use of the name punica by various authors for populations of M. ornata in a number of different countries, Melitaea punica is entirely confined to North Africa, where it is restricted to Morocco and Algeria; there have been no reports from Tunisia (see Appendix: Note 3).

Synonymic list

Melitaea phoebe abbas Gross & Ebert, 1975 Note 7 [Type Locality (TL): 50 km. NW Ardkan, Tange Sorkh, Fars, Iran, 2250 m, 12–15.vi.1975].

Melitaea ornata adversaria Korb, Stradomsky & Kuznetsov, 2015 Note 8 [TL: Kirghizia, Kirghiz Mts., Ala-Too settlement vicinity, 1100–1200 m].

Melitaea phoebe var. aetherea Eversmann, 1851 Note 9 [TL: Russia ‘au Sud qu’au Nord’].

Melitaea phoebe aethereaeformis Verity, 1919 Note 10 [TL: central Italy].

Melitaea phoebe alatauica Wagner, 1913 Note 11 [TL: Issyk-kul, Ili mountains, Kazakhstan].

Melitaea phoebe ab. albina Verity, 1904 Note 12 [TL: Lucca, Italy]

Melitaea phoebe allophylus Rütimeyer, 1942 Note 13 [TL: Porté, Pyrénées Orientales, France].

Melitaea phoebe almana Gaede, 1930 Note 14 [TL: Elman Dagh N Syria]

Melitaea phoebe alternans Seitz, 1909 Note 15 [TL: Zermatt, Switzerland].

Melitaea phoebe Knoch var. amanica Rebel, 1917 Note 16 [TL: Kushdjula, Taurus Mountains and Das Dagh, Amanus Mountains, Turkey].

Melitaea phoebe rovia autumnalis Fruhstorfer, 1919 Note 17 [TL: between Brione & Contra, Italy].

Melitaea phoebe ab. baccata Delahaye, 1909 Note 18 [TL: Saint-Barthélemy, Maine-et-Loire, France].

Melitaea phoebe Knoch rassa bethune-bakeri de Sagarra, 1926 Note 19 [TL: Sierra Nevada, Spain].

Melitaea phoebe Knoch sbsp. n. canellina Stauder, 1922 Note 20 [TL: vicinity of Innsbruck, Austria].

Melitaea phoebe capreola Varga, 1967 Note 21 [TL: Podalia, Kiverci, Ukraine].

Melitaea phoebe var. caucasica Staudinger, 1870 Note 22 [TL: “Kindermann ganz ähnliche Stücke im Caucasus fing (?-Helenendorf; Kindermann leg.)”].

Melitaea phoebe caucasicola Verity, 1919 Note 23 [TL: “Kindermann ganz ähnliche Stücke im Caucasus fing (?-Helenendorf; Kindermann leg.)”].

Melitaea phoebe changaica Seitz, 1909 Note 24 [TL: Changai Mountains, Mongolia].

Melitaea phoebe ab. cinxioides Muschamp, 1905 Note 25 [TL: Switzerland].

Melitaea phoebe Knoch ab. confusa Joannis, 1908 Note 26 [TL: Vannes, France].

P.[apilio] N. Phal. [Nymphalis Phaleratus] corythallia Esper, [1781] Note 27 [TL: France (environs of Paris?)]

Melitaea phoebe crassenigra Verity, 1928 Note 28 [TL: Rozier, Gironde, France].

Melitaea phoebe form deleta Verity, 1919 Note 29 [TL: Tuscany, Italy].

Melitaea phoebe Knoch dorae Graves, 1925 Note 30 [TL: Nabatea, Petra, Jordan].

Melitaea phoebe tusca emipauper Verity, 1919 Notes 31 & 96 [TL: Vallombrosa, Tuscany, Italy].

Melitaea phoebe emipunica Verity, 1919 Note 32 [TL: Palermo district, Sicily, Italy].

Melitaea phoebe mod. enoch Higgins, 1941 Note 33 [TL: Askhabad, Turkmenistan].

Melitaea phoebe occitanica f. estrela Higgins, 1941 Note 34 [TL: Sierra de Estrela, Portugal]

Melitaea phoebe Knoch ab. fasciata Galvagni, 1934 Note 35 [TL: ‘Austria Inferior’].

Melitaea phoebe Knoch rassa occitanica Staudinger 2-gen. francescoi de Sagarra, 1926 Notes 5 & 36 [TL: between St. Pere & Vilamajor, Catalonia, Spain, July/August].

Melitaea phoebe gaisericus Hemming, 1941 Note 37 [TL: Mogador (=Essaouira), Morocco].

Melitaea phoebe galliaemontium Verity, 1928 Note 38 [TL: Mont Dore, Puy de Dome, France].

Melitaea phoebe gerinia Fruhstorfer, 1917 Note 39 [TL: Lisbon, Portugal].

Melitaea phoebe ab. geyeri Aigner-Abafi, 1906 Note 40 [TL: Szaár (Komitat Fejér), Hungary].

Melitaea phoebe guevara Fruhstorfer, 1917 Note 41 [TL: Castile, Spain].

Melitaea phoebe Knoch ab. gürtleri Joukl, 1908 Note 42 [TL: Plitvička Jezera, Croatia]

Melitaea phoebe occitanica f. juliae Molina & Ocete, 1986 Note 43 [TL: Loma de la Amoladera, Guadalcanal (Seville), Spain]

Melitaea phoebe koios Fruhstorfer, 1908 Note 44 [TL: Klausen, Switzerland].

Melitaea phoebe kovacsi Varga, 1967 Note 45 [TL: Budakeszi, Hungary].

Melitaea phoebe leechi Rothschild, 1917 Note 46 [TL: Mogador (= Essaouira), Morocco].

Melitaea phoebe lokris Fruhstorfer, 1908 Note 47 [TL: Saratov, Russia].

Melitaea phoebe malvida Gaede, 1930 Note 48 [TL: Meklen Pass, Bosnia].

Melitaea phoebe mandarina Seitz, 1909 Note 49 [TL: Mongolia].

Melitaea phoebe var. melanina Bonaparte, 1831 Note 50 [TL: Monti Subiaco (= Livata), near Santa Scolastica, Arbruzzo, Italy.

Melitaea phoebe minoa Fruhstorfer, 1917 Note 51 [TL: Engadin, Switzerland].

Melitaea phoebe ab. minor Wheeler, 1903 Note 52 [TL: Switzerland].

Melitaea phoebe monilata Verity, 1919 Note 53 [TL: Wallis (= Valais), Switzerland].

Melitaea phoebe monilataeformis Verity, 1919 Notes 54 & 96 [TL: Tuscany, Italy].

Melitaea phoebe narenta Fruhstorfer, 1917 Note 55 [TL: Jablanica, Herzegovina].

Melitaea phoebe nigroalternans Verity, 1919 Note 56 [TL: Mont Cenis, French/Italian border].

Melitaea phoebe nigrogygia Verity, 1939 Note 57 [TL: Abbazia = Opatija, Istria, Croatia].

Melitaea phoebe mod. nimbula Higgins, 1941 Note 58 [TL: Espinama, Picos de Europa, Cantabria, Spain].

Melitaea phoebe occitanica Staudinger, 1871 Note 5 [TL: Andalusia, Spain].

Melitaea phoebe ogygia Fruhstorfer, 1907 Note 59 [TL: Island of Poros, Greece].

Melitaea phoebe ornata Christoph, 1893 Notes 2 & 6 [TL: Circa ‘Guberli’, promontorium uralensium australium (Guberlya, Orenburg Province, Russian Federation)].

Melitaea phoebe Schiff. ornatiformis (gen. aestiva) de Sagarra, 1930 Note 60 [TL: Villacabras, Cuenca, Spain].

Melitaea phoebe ottonis Fruhstorfer, 1917 Note 61 [TL: “Kindermann ganz ähnliche Stücke im Caucasus fing (?-Helenendorf; Kindermann leg.)”].

P.[apilio] NP Paedotrophos Bergsträsser, 1780 Note 62 [TL: Hanau-Münzenberg, Germany]

Melitaea phoebe subsp. parascotosia Collier, 1933 Note 63 [TL: Sutschan, Russian Federation].

Melitaea phoebe ab. parva Gerhard, 1882 Note 64 [TL: Fünfkirchen (= Pecs), Hungary].

Melitaea phoebe var. parva Caradja, 1895 Note 65 [TL: Bucharest, Romania].

Melitaea phoebe pauper Verity, 1919 Notes 66 & 96 [TL: Florence, Italy].

P.[apilio] phoebe Denis & Schiffermüller, 1775 Notes 1, 4 & 6 [TL: environs of Vienna, Austria].

Melitaea phoebe phoebina Turati, 1919 Note 67 [TL: Aspromonte Mountains, Calabria, Italy].

Melitaea phoebe postnarenta Verity, 1939 Note 68 [TL: St. Dionisio, Mt. Olympos, Greece].

Melitaea phoebe postogygia Verity, 1939 Note 69 [TL: Salonica (= Thessalonica), Greece.

Melitaea phoebe virgilia postvirgilia Verity, 1950 Notes 70 & 100 [TL: Vence, Alpes-Maritimes, France].

Melitaea phoebe pseudosibina Alberti, 1969 Note 71 [TL: Mt. Elbrus, Itkol, Kabardino-Balkaria, Russia].

Melitaea phoebe punica Oberthür, 1876 Notes 3 & 6 [TL: Tazoult-Lambèze (Lambessa), Algeria].

Melitaea phoebe punica-powelli Oberthür, 1915 Note 72 [TL: Algeria].

Melitaea phoebe forma punicata Ragusa, 1919 Note 73 [TL: Sicily, Italy].

Melitaea ornata reliquiae Korb, Stradomsky & Kuznetsov, 2015 Note 74 [TL: Russia, Volgograd Province, Olkhovsky distr., Kamenny Brod].

Melitaea phoebe rostagnoi Turati, 1920 Notes 75 & 96 [TL: Rome, Italy].

Melitaea phoebe rovia Fruhstorfer, 1919 Note 76 [TL: Monte Generoso, Maroggia, Tessin, Switzerland].

Melitaea phoebe f. rubialesi Gómez Bustillo, 1973 Note 77 [TL: Loeches (Madrid), Spain]

Melitaea phoebe forma rubrofasciata Gušić, 1922 Note 78 [TL: Podsused, nr. Zagreb, Croatia].

Melitaea phoebe Knoch sarvistana Wiltshire, 1941 Note 79 [TL: Sarvistan, SE of Shiraz salt lake, Iran].

Melitaea phoebe var. saturata Staudinger, 1892 Note 80 [TL: Kentai Mountains, Mongolia].

Melitaea phoebe ab. seminigra Delahaye, 1909 Note 81 [TL: Pignerolles, Maine-et-Loire, France].

Melitaea phoebe Knoch var. sextilis Jachontov, 1909 Note 82 [TL: Zheleznovodsk, Stavropol Krai, Russian Federation (North Caucasus)].

Melitaea phoebe Knoch var. sibina Alphéraky, 1881 Note 83 [TL: Kuldjà, Ili Valley, China].

Melitaea phoebe rostagnoi ab. sterlineata Turati, 1920 Note 84 [TL: Monte Autore (Province of Rome), Italy].

Melitaea phoebe streltzovi Kolesnichenko & Yakovlev, 2004 Note 85 [TL: Western Mongolia, Hovd aimak, 30 km north-northwest from Bulgan somon, junction of Bajan-Gol and Bulgan-Gol rivers, 1500 m 11–13 August 2003].

Melitaea phoebe subcorythallia Verity, 1928 Note 86 [TL: Auzay, Vendée, France].

Melitaea phoebe suboccitanica Verity, 1928 Note 87 [TL: Auzay, Vendée, France].

Melitaea phoebe subtusca Verity, 1952 Notes 88 & 96 [TL: Nans-les-Pins, St. Baume, Var, France].

Melitaea phoebe sylleion Fruhstorfer, 1917 Note 89 [TL: Cogne, Piedmont, Italy].

Melitaea phoebe tatara Krulikovsky, 1891 Note 90 [TL: Casanum = Kazan or Casan, Republic of Tatarstan, Russian Federation].

Melitaea phoebe telona Fruhstorfer, 1907 Note 91 [TL: Jerusalem, Palästina (Israel)].

Melitaea phoebe forma totila Stauder, 1914 Note 92 [TL: Monte Cocuzzo, Consenza, Calabria, Italy].

Papilio tremulae Piller & Mitterpacher, 1783 Note 93 [TL: between Drau & Sawe, Croatia].

Melitaea phoebe tungana Seitz, 1909 Note 94 [TL: “Sajan District”, Russian Federation].

Melitaea phoebe tungusa Herz, 1899 Note 95 [TL: Witim & Vilui mountains, Siberia, Russian Federation].

Melitaea phoebe tusca Verity, 1919 Note 96 [TL: Tuscany, Italy].

Melitaea phoebe var. occitanica ab. uclensis Melcón, 1910 Note 97 [TL: Uclo, Cuenca, Spain].

Melitaea phoebe ufensis Krulikovsky, 1902 Note 98 [replacement name for uralensis Note 99].

Melitaea phoebe uralensis Krulikovsky, 1897 Note 99 [TL: district of Ufa, Russia].

Melitaea phoebe virgilia Fruhstorfer, 1917 Note 100 [TL: Alpes Maritimes, France].

Melitaea phoebe wagneri Wnukowsky, 1929 Notes 11 & 101 [replacement name for alatauica Wagner].

Melitaea phoebe scotosia yagii Nire, 1917 Note 102 [TL: c. 5 km west of Mt. Asama, Shinano Province, Japan].

Melitaea zagrosi Tóth & Varga, 2011 Note 103 [TL: Zagros Mountains, Iran].

Acknowledgements

James Pateman (Tangmere, UK) is thanked for his dissections and comments regarding genitalia of specimens of M. phoebe and M. ornata from Montenegro and Russia. Assistance from Lazaros Pamperis (Larissa, Greece) in providing information on the presently known distribution of M. phoebe on mainland Greece was much appreciated. John Coutsis (Athens, Greece) is thanked for discussion on the origin of the type locality of the name nigrogygia. Dusan Zitnan (Bratislava, Slovakia) is thanked for translating the article by Joukl from the original Czech. Provision of a copy of the Delahaye (1906) supplement by Eric Drouet (Gap, France) was greatly appreciated, as was provision of Nire’s 1917 paper by Akio Masui (Japan).

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Appendix

Note 1. Melitaea phoebe (Denis & Schiffermüller, 1775) [Type Locality (TL): environs of Vienna, Austria]: type material lost; neotype ♂ designated by Tennent and Russell (2010), reared from larva collected from near Vienna, Austria. An often double or even triple brooded species.

Note 2. ornata: (Melitaea ornata) Christoph, 1893 [TL: Circa ‘Guberli’, promontorium uralensium australium (near Guberlya, Orenburg Province, Russian Federation)]: this taxon was first recognised as a species distinct from phoebe by Tóth and Varga (2011), based on morphometric measurements of male and female genitalia. It was discovered in the Volgograd region at Ilovlya by Tuzov and Churkin (2000: 73, pl. 46, figs 7–9 & 15–17) who wrongly used the name M. (phoebe) punica; Kuznetsov and Stradomsky (2010) subsequently used the name Melitaea telona. Kuznetsov (2011) provided details of the biology of this taxon and Russell and Kuznetsov (2012: figs 1–3) demonstrated that larvae from the Volgograd region had red-brown heads. This character is diagnostic for M. ornata (within the Melitaea phoebe species group – M. cinxia larvae also have red-brown heads).

The colourful adults were illustrated by Higgins (1941: pl. 14, fig. 10), Gorbunov and Kosterin (2007: 2: 84, figs 197, 198) and van Oorschot and Coutsis (2014: pl. 12: figs 20, 21 & pl. 13: figs 2, 3). M. ornata contrasts with the less colourful M. phoebe flying at the same localities in the southern Ural Mountains (present authors, pers. obs.). Tshikolovets (2011: 498) and Tshikolovets et al. (2014: 319) recognised ornata as a distinct species. The fact that van Oorschot and Coutsis (2014: 60) placed ornata as a synonym of nominotypical phoebe is in part what prompted the present paper. The first author has reared many hundreds of specimens of both phoebe and ornata from many localities in Europe and both authors have seen adults of the two species (sympatric but not synchronic) flying in the Urals. There can be no doubt whatever that Melitaea phoebe and Melitaea ornata (=telona: see Note 91) are distinct species, with different early stages, voltinism (M. ornata is invariably single brooded M. phoebe often double or triple brooded) and they often have different host-plants.

Recognition of ornata as a distinct species paved the way for the realisation that what had recently been referred to as “emipunica” (Russell et al. 2005), “ogygia” (Varga et al. 2005) and “telona” (Kuznetsov and Stradomsky, 2010) all represented the same species (i.e. ornata).

Note 3. punica: (Melitaea punica) Oberthür, 1876 [TL: Tazoult-Lambèze (Lambessa), Algeria]: This species was described by Oberthür (1876: 25) as a subspecies of M. phoebe, but Oberthür himself subsequently raised it to the status of a distinct species (Oberthür 1914: 102). It is now recognised as a distinct species by most modern authors. Following the unfortunate introduction of a quadrinomen “M. (phoebe) punica telona” (this actually encompassed three distinct species: phoebe, punica and ornata [as telona]) by Hesselbarth et al. (1995: 1030), the status of punica became very confused. It was repeated in that form by Koçak (2000: 9), and a number of authors (e.g. Koçak and Seven 1998: 4) used the combination “Melitaea punica telona”. Nazari (2003) placed all the taxa mentioned (including telona but not punica) as synonyms of M. phoebe, with the rather unhelpful note: “For further synonymy see Higgins (1941: 338–343)”.

Subsequently, and presumably as a direct result of the action by Hesselbarth et al. (1995: 1030), the name punica was frequently wrongly associated with the name telona Fruhstorfer, 1908 (see also Note 2). For example Abadjiev (2000, 2001: 271), Tuzov and Churkin (2000: 73, pl. 46, figs 7–9, 15–17), Rákosy and Varga (2001), Gorbunov and Kosterin (2003 2: 84) and more recently Baytaş (2007: 128), Székely (2008: 175), Aghababyan (2012: 13), Hüseyinoğlu and Akyol (2013: 11 & 14) and Hüseyinoğlu (2013: 1293), all used the combination ‘Melitaea (Cinclidia) (phoebe) punica’ for the taxon ornata. This confusion was undoubtedly brought about by the fact that the underside hindwing pattern (particularly in the submarginal area) of M. punica (cf. Russell et al. 2006: figs 12–26) is very similar to those non-phoebe specimens from Europe and Turkey. This was clearly demonstrated by Russell and Pamperis (2011: 140–142 & figs 3–8; 2012) and Russell and Pateman (2012: figs 4–7). Other authors simply used the name punica for the species which was not M. phoebe s.s.: e.g. Leraut (1999: 173), who gave the distribution of “C[inclidia] punica” (i.e. Melitaea punica) as Italy to Turkey and Jordan, with no mention of North Africa, the TL of punica and the only place where M. punica is actually known to occur.

More recently, Tóth and Varga (2011) and van Oorschot and Coutsis (2014: 66) separated punica from phoebe on the basis of differences in the male genitalia, and this was followed by Tshikolovets (2011: 497). Collectively, the published literature during the last two decades has created substantial confusion which, insofar as it affects M. punica, is clarified here: Melitaea punica is confined to North Africa; it occurs from the Atlantic coast of Morocco throughout the Atlas and Rif Mountains to eastern Algeria but apparently not into Tunisia (Tennent 1996: 52). The larva is very similar to that of M. phoebe occitanica (Note 5) (Russell et al. 2006: figs 1–4 & 6); however, punica butterflies in North Africa are quite different in appearance to phoebe occitanica in Spain (cf. Higgins 1941: pl. 14, figs 6 & 11; Russell et al. 2006: figs 8–26; Tolman and Lewington 2008: 203).

Note 4. Melitaea phoebe phoebe: The body of the final instar larva of M. phoebe phoebe is black, including the head carapace, with black or orange tubercles and white spots spaced around each segment; these spots usually coalesce on each side to form an often prominent lateral white line (see Table 1; also Bodi 1985: plate XI, fig. 92; Lafranchis 2000: 391, fig.; Russell et al. 2007: 159, fig. 14; Lafranchis 2007a: 41, fig. 13; Lafranchis 2008: 6 (fig.); Tennent and Russell 2010: 151, fig. 9). Its distribution ranges from the Ural Mountains to c. 60º N., through the Caucasus, south to Iraq, Iran and Lebanon, reaching its southern limit in northern Israel (Benyamini pers. comm.), westwards through Turkey, the Balkans, Hungary, Austria, southern Germany and the alpine and sub-alpine regions of France, Switzerland and Italy.

Some of the name bearing types originate from the eastern Palaearctic. For the sake of completeness these have been included. They are synonymised with nominotypical phoebe due to the fact that Kosterin (see Korshunov and Gorbunov 1995) described a final instar larva of M. phoebe from near Zabaikalye (south-eastern Russia) as follows: “white with fine black reticulate ornament, so that looks grey; this ornament fuses into a black line along the back and a more diffuse line on either side (between 2nd and 3rd row of false spines from beneath); a white stripe (without ornament) goes through 2nd row beneath false spine row. Thoracic legs and ventral prolegs yellowish-grey; head greyish-black, set with tiny black hairs”. This description precisely matches that of the final instar larvae of the European populations of nominotypical phoebe. Adult butterflies are very variable, and we are unable to separate populations of phoebe s.s. in western Europe to the Urals into distinct races (subspecies).

Note 5. Melitaea phoebe occitanica Staudinger, 1871 [TL: Andalusia, Spain]: the Type Locality of this subspecies was given originally by Staudinger as “It” (= Italy?) but it is now generally accepted that this was an error (recte ‘Iberia’: Higgins 1941: 336); Verity (1928: 163) limited the Type Locality to Barcelona, Spain, and this was accepted by van Oorschot and Coutsis (2014: 60). Previously, however, Fruhstorfer (1916: 82 (A) (2): 1) was of the opinion that the source of the occitanica phenotype was Andalusia and Higgins (1941: 336) considered that this should stand, based on the original description by Staudinger, who did not specify a ‘Type’ but labelled the series upon which the description was based with the word ‘original’ (Higgins 1941: 336). The body of the final instar larva of M. phoebe occitanica is black, including the head, with obscure white spots on the body, black tubercles dorsally and a row of orange tubercles with orange hairs laterally, which form an obvious orange lateral line (Lafranchis 2000: 388, fig.; Maravalhas 2003: 281, fig.; Russell et al. 2007: 159, fig. 13; Lafranchis et al. 2015: 464–467, figs), in contrast to the white lateral line of nominotypical phoebe (see Table 1, and Note 4).

This subspecies has also been separated from nominotypical phoebe using the results of enzyme electrophoresis by Pelz (1995: 57), who was of the opinion that genetic differences were sufficiently significant for occitanica to be considered as a “semispecies”. This subspecies is distributed from the Iberian Peninsula eastwards through France and peninsular Italy as far south as northern Calabria (Russell pers. obs.); it has also been found in Istria, Croatia (Russell and Pateman 2013a: 47, fig. 6).

Tshikolovets (2011: 496) suggested that the distribution of this subspecies included northern Greece, the southern and eastern parts of the Balkans, western Turkey and Ukraine. The present authors do not agree and consider that these areas are occupied by nominotypical phoebe; larvae from Romania, for example, are clearly of the ‘white lateral stripe form’ associated with nominotypical phoebe (Russell et al. 2007: 159, fig. 13). Where the two subspecies meet, for example in Var, France and Istria, Croatia, the larvae can be intermediate in form, as one might expect (Russell and Pateman 2013: 47, figs 8, 9). The colourful adult has been illustrated by many authors, including Higgins (1941: pl. 14, fig. 11), Manley and Allcard (1970: plate 10, figs 1–7) and Lewington in Tolman and Lewington (1997: plate 50).

Note 6. The species phoebe, punica, ornata: despite a series of articles (e.g. Russell et al. 2005, 2006, 2007), Tolman and Lewington (2008: 202–203) recognised only one species M. phoebe. However, all three species were separated using DNA sequences by Lenevue et al. (2009) and Tóth et al. 2014. Recognition of these distinct species is now accepted by most recent authors (Tshikolovets 2011: 496–499; Tóth and Varga 2011; van Oorschot and Coutsis 2014: 60–64 & 66), although not necessarily using correct species and/or subspecies names in the correct combinations. The raison d’être for this paper is to resolve this nomenclatural muddle.

Note 7. abbas Gross & Ebert, 1975: 44, fig. 61: synonym of Melitaea ornata. Tshikolovets et al. (2014: 320, map) gave this taxon subspecific status of M. ornata and stated that it was found in west and south Iran. Their figures (Plate LX: figs 7, 8, 10, 11 & 12) depicted specimens which appear to have spatulate antennae and black arched submarginal underside hindwing markings not touching the intervening veins, both features typical of M. ornata (see Table 1). Subsequently, van Oorschot and Coutsis (2014) figured 5 specimens (plate 13: figs 8, 11, 12, 13 & 16), all from western Iran, of M. ornata (as M. telona).

Note 8. Melitaea ornata adversaria Korb, Stradomsky & Kuznetsov, 2015: 142 & plate VI: tentative synonym of Melitaea ornata. This material has been classified as both Melitaea phoebe saturata (Korb 2011: 158: see Note 80) and Melitaea ornata adversaria (Korb et al. 2015). The latter was based on molecular analysis of the preserved specimens and we Note that Korb et al. (2015: 142) considered that M. phoebe was not present in the Kyrghyz Mountains. The flight period was given (Korb 2011: 158) as May–July, at elevations between 500 and 2000 m; we consider that July is likely to be beyond the flight time of M. ornata and that larvae would be in diapause by the end of June. So far as we are aware, larvae of the Melitaea populations in this region have not been reported; our synonymy is thus tentative, pending further data.

Note 9. aetherea Eversmann, 1851: 5: 73 and plate IX: figs 5, 6: synonym of nominotypical phoebe. Synonymised with M. phoebe by Higgins (1941), and followed by van Oorschot and Coutsis (2014: 60). A large but weakly marked form (Higgins 1941: 338, plate 14: fig. 9). Korshunov and Gorbunov (1995: species 174) gave a very full description of the larva of this form, which clearly associated it with nominotypical phoebe. Tshikolovets (2011: 497) used this name at subspecies rank.

Note 10. aethereaeformis Verity, 1919: 183: synonym of phoebe occitanica. Specimens from central Italy which were similar in appearance to aetherea Eversmann, 1851 (Higgins 1941: 338) were due to its geographical location placed with occitanica. Overlooked by van Oorschot and Coutsis (2014).

Note 11. alatauica Wagner, 1913, vol. 2: 89 (fig.): Junior primary homonym of M. parthenie alatauica Staudinger, 1881, and presumed synonym of nominotypical phoebe. The authors consider that this name is most probably related to M. phoebe since it occurs in the eastern Palaearctic outside the presently known eastern limit of the distribution of M. ornata (see introduction). Placed with M. phoebe by van Oorschot and Coutsis (2014: 60), who suggested that it could be synonymous with M. sibina Alphéraky, 1881 (see Note 83).

Note 12. albina Verity, 1904: 54: we cannot place this form with either M. phoebe or M. ornata. An aberrant individual having the ground colour of the right hindwing yellowish-white (Higgins 1941: 339); both species may occur in the Lucca region of Italy.

Note 13. allophylus Rütimeyer, 1942: 438: synonym of phoebe occitanica. Higgins (1955: 118) recognised this form as “leading to occitanica Staudinger” and suggested that it should be synonymised with M. corythallia Esper, 1781 (i.e. phoebe occitanica, see Note 27).

Note 14. almana Gaede, 1930: 208: probable synonym of Melitaea ornata. This name, attributed to Rebel, appears to have been first published by Gaede under M. phoebe (in Seitz, Supplement). Neither Higgins (1941: 339) nor the present authors were successful in their efforts to find an original Rebel reference, and as a result it is provisionally treated as a Gaede manuscript name. Gaede stated that it was a pale race from Asia Minor similar to M. telona (i.e. ornata). Hesselbarth et al. (1995: 1031) referred to Graves (1925: 101), who stated that this form came from Elma Dagh, Syria. They suggested that it may have been a misspelling of amanica Rebel (see Note 16) and synonymised it with Melitaea punica telona (i.e. ornata), although it is Noted that Gaede treated both names separately. Not mentioned by van Oorschot and Coutsis (2014).

Note 15. alternans Seitz, 1909: 216: synonym of nominotypical phoebe. A large brightly coloured subalpine form; not figured by either Seitz (1909) or Higgins (1941) but figured by Tolman and Lewington (1997: plate 50; 2008: 203 [same painting]); Higgins (1941: 339) suggested that it was ‘proceeding to occitanica Staudinger’ but only because of its brighter colouring, which is typical of both Alpine and Spanish specimens. See also monilata (Note 53).

Note 16. amanica Rebel, 1917: 252: synonym of Melitaea ornata. Tshikolovets and Nekrutenko (2012: 295) synonymised this form with telona, placing the latter as a subspecies of M. ornata, and recorded its distribution as the Lesser Caucasus, Djavakheti-Armenian plateau and Talysh. The form is univoltine, with a flight period of May (sometimes late April) – June. Tuzov et al. (2000: plate 46: figs 7–9) figured three specimens in colour with the legends: “Melitaea (phoebe) punica amanica Rebel”, two from Armenia, Azavan and one from Azerbaijan, Talysh Mts, Zuvand Plateau, Gosmalyan, 1500 m, 4.vi.1981. Antennal clubs of these specimens appear short and the hindwing underside markings in the submarginal area appear similar to those of M. ornata. Van Oorschot and Coutsis (2014: plate 13, fig. 7) figured in colour a specimen from Armenia, Vedi, vicinity of Chosrov, 27.v.1974, under the name M. telona (i.e. ornata), which appears from its underside hindwing markings to be correct.

Note 17. autumnalis Fruhstorfer, 1919: 169: synonym of nominotypical phoebe. The second generation form of rovia Fruhstorfer, 1919 (see Note 76). Bernardi and de Lesse (1951: 141) identified a holotype for autumnalis (as automnalis).

Note 18. baccata Delahaye, 1909: 10: aberration of phoebe occitanica. The supplement in which this name was published was not available to Higgins (1941: 339), but was kindly supplied to the authors by Eric Drouet. The name refers to an aberrant female specimen which was taken in August at Saint-Barthélemy, Maine-et-Loire in west-central France and thus outside the known ranges of both nominotypical phoebe and M. ornata. Not mentioned by any recent author.

Note 19. bethunebakeri de Sagarra, 1926: 130: synonym of phoebe occitanica. Higgins (1941: 339) correctly considered it synonymous with occitanica Staudinger 1871. Not mentioned by van Oorschot and Coutsis (2014).

Note 20. canellina Stauder, 1922: 18: synonym of nominotypical phoebe. Higgins (1941: 339) suggested this was synonymous with minoa Fruhstorfer, 1917 (see Note 51); the TL places it outside the known ranges of both phoebe occitanica and ornata but within the distribution of nominotypical phoebe. Overlooked by van Oorschot and Coutsis (2014).

Note 21. capreola Varga, 1967: 131: synonym of Melitaea ornata. Varga described this as a subspecies of M. phoebe, but subsequently (Tóth and Varga 2011) placed it with M. ornata; van Oorschot and Coutsis (2014: 63) placed it with M. telona (i.e. ornata).

Note 22. caucasica Staudinger, 1870: 59, Taf. 1 fig. 2: synonym of nominotypical phoebe, but name preoccupied by M. didyma caucasica Staudinger, 1861; see ottonis Fruhstorfer, 1916 (a replacement name for caucasica: Note 61), and caucasicola Verity, 1919 (Note 23), a later replacement name. A lectotype ♀ and a paralectotype ♂ were designated by Nekrutenko (Hesselbarth et al. 2: 1028) from the Staudinger collection, housed at Zoologisches Museum der Humboldt Universität, Berlin.

Note 23. caucasicola Verity, 1919: 184: a replacement name for caucasica Staudinger, 1870 (see Note 22); a junior subjective synonym of ottonis Fruhstorfer, 1916 (see Note 61).

Note 24. changaica Seitz, 1909: 217: synonym (provisional) of nominotypical phoebe. Occurs in the eastern Palaearctic, further east than the presently known eastern limit of the distribution of M. ornata. Kosterin figured a final instar larva of this taxon from 10 km NNW of the village of Tasyrkhoi S Chita region (Dahuria), Transbaikalia, Siberia, Russia, 19.vi.1995. Its black head carapace confirms probable synonymy with M. phoebe.

Note 25. cinxioides Muschamp, 1905: 69 (fig.): aberrational form of nominotypical phoebe. Its origin in Switzerland is outside the distributional areas of both phoebe occitanica and M. ornata. An aberrant form with black spots in the submarginal brown spots of the hindwing upperside, resembling M. cinxia. This recurrent aberration is known to occur almost anywhere (pers. obs.). Placed with M. phoebe by Higgins (1941: 339). Not mentioned by any recent authors.

Note 26. confusa Joannis, 1908: 45: synonym of phoebe occitanica. An aberrant ♂ form in which the upperside forewings are more reddish with the black markings reduced, the transverse black lines in the discal region are nearly obliterated and the hindwings are dark basally. The underside forewings have similar markings but the hindwings are yellowish white with enlarged dark markings. Higgins (1941: 339) attributed this name to Oberthür but with Joannis’ reference, and he did not correct this in his errata (Higgins 1944). The TL of Brittany, northwest France, places it outside the known ranges of nominotypical phoebe and M. ornata. Not mentioned by any recent authors.

Note 27. corythallia Esper, [1781]: 65, 67, Taf. 61, figs 4, 5: synonym of phoebe occitanica. Verity (1928: 163) was of the opinion that occitanica Staudinger (see Note 5) should be placed as a synonym of corythallia Esper, on the basis that he believed the specimens representing corythallia originated from the Iberian Peninsula. Higgins (1941: 336) disagreed with this course of action and showed that Verity’s assumption was incorrect, as Esper ([1781]: 67), stated that they were the original specimens of Geoffroy’s Papilio cinxia var. B, which were from France (Higgins 1941: 336). Whether the origin of the specimens of corythallia were from France or Spain is unimportant because the same subspecies of M. phoebe (i.e. occitanica) occurs in both countries. Hesselbarth et al. (1995: 1028) and van Oorschot and Coutsis (2014: 60), synonymised this name with M. phoebe. Although the name corythallia predates occitanica, type material of the former appears to be lost (Hesselbarth et al. 1995: 1028) and the name occitanica has been used extensively by authors in referring to phoebe populations from the Iberian Peninsula. The present authors have followed this course of action.

Note 28. crassenigra Verity, 1928: 162: synonym of phoebe occitanica. An occitanica form with heavy discal spotting from southwest France (Higgins 1941: 339).

Note 29. deleta Verity, 1919: 184: aberration of (presumably) phoebe occitanica. Aberrant female of form tusca (see Note 96) with almost all the black markings obliterated (Higgins 1941: 339). Larvae reared from populations of M. phoebe from peninsular Italy have, so far as the authors are aware, all been of the occitanica form (see Note 5).

Note 30. dorae Graves, 1925: 100: synonym of Melitaea ornata. Graves (1925: 103–106) gave a two page description of this form, and a table of “Index of Nigrescence of M. phoebe races (upperside)”, which demonstrated that it was paler than either telona or ogygia. Higgins (1941: 339) paraphrased this description as “small and pale, with the black markings fine and partly obsolete”; this is typical of phenotypes in xerothermic biotopes. Hesselbarth et al. (1995: 1031) synonymised this name with ‘M. punica telona’ (i.e. ornata) and Tshikolovets (2011: 499) with ‘Melitaea ornata telona’ (i.e. ornata). Van Oorschot and Coutsis (2014: 63) synonymised this with M. telona (i.e. ornata) and illustrated (van Oorschot and Coutsis 2014: plate 13, fig. 6) a specimen from Wadi Zarqa, Jordan, 400 m, the underside hindwing pattern and spatulate antennae of which suggest synonymy with ornata.

Note 31. emipauper Verity, 1919: 184: synonym of phoebe occitanica. Described by Verity as a medium sized, summer brood form of tusca Verity, 1919 (Higgins 1941: 340) (see Note 96).

Note 32. emipunica Verity, 1919: 184: synonym of Melitaea ornata. This name was used by Russell et al. (2005) when the species was first identified as being distinct from M. phoebe by the red-brown head colour of the stage L4 to the final instar larvae, reared from a female taken at Montagna Longa, within the Type Locality (i.e. Sicily). This was afforded subspecific status by Tshikolovets (2011), with a distribution given as SE France (Var), Sicily and S Italy (Calabria, Basilicata, Campania). It was synonymised with M. telona (i.e. ornata) by van Oorschot and Coutsis (2014: 63). Brief use of emipunica by Russell et al. (2005) and of ogygia by Varga et al. (2005) was before the wide distribution of M. ornata was fully appreciated, and was (in part) the cause of ensuing confusion.

Note 33. enoch Higgins, 1941: 337: synonym of Melitaea ornata. Figured by Higgins (1941: plate 14, fig. 4), who gave other locations for this form: Arwas and Achal Tekke, 2000 m, July, and Jablonowka from the same region (Transcaspia). He placed it with M. phoebe occitanica but suggested that this placement was due to the colour contrast of the wings being similar to, but not quite so strongly marked as, those of ‘Spanish occitanica’. Higgins further noted a slight difference in male genitalia and suggested the possibility that enoch should be ranked as a subspecies of M. phoebe. Tshikolovets et al. (2014: 319 and plate LX: figs 6 and 9) placed enoch as a subspecies of M. ornata. Van Oorschot and Coutsis (2014: 63) placed it as a synonym of M. telona (i.e. ornata).

Note 34. estrela Higgins, 1941: 337: phoebe occitanica. Said by Higgins (1941: 337) to be: “very bright … labelled estrela Romei, but I cannot trace a reference to a description, and do not know whether the name was ever published validly …”; the present authors have also failed to find a published reference by Romei, and place the name as a nomen nudum.

Note 35. fasciata Galvagni, 1934: 2: an aberration of nominotypical phoebe. This extreme aberration has the upper surface of the wings almost black with the forewing discal macules radially elongated into a fascia. The specimen was taken on 6.viii.1933 near Vienna; its origin places it with nominotypical phoebe. The name has been used by a number of authors to describe specimens in which the black markings coalesce to form fasciae; for example Wiltshire (1946: 26; plate 3, fig. d) used it to describe a specimen of M. phoebe from Shiraz, Fars, SW Iran, suggesting it was similar to “mod. or ssp. telona” (= ornata).

Note 36. francescoi de Sagarra, 1926: 130: synonym of phoebe occitanica. A name raised for specimens of the second brood of occitanica Staudinger, 1871, flying in July/August (Higgins 1941: 340) (see also Note 5).

Note 37. gaisericus Hemming, 1941: 207: synonym of Melitaea punica. A replacement name for leechi Rothschild (see Note 46) (Higgins 1941: 340); Higgins (1941: pl. 15, fig. 8) figured an example from Azrou, Morocco. Synonymised with M. punica by van Oorschot and Coutsis (2014: 66).

Note 38. galliaemontium Verity, 1928: 162: synonym of phoebe occitanica. A name raised for small, second brood specimens (Higgins 1941: 340) from France. Overlooked by van Oorschot and Coutsis (2014).

Note 39. gerinia Fruhstorfer, 1917: 1: synonym of phoebe occitanica. This form is more uniform in colour than the contrasting highly coloured form found in Spain (Higgins 1944: 340) (see Note 15). In raising the name gerinia, Fruhstorfer (1917: 1–2) did not refer to specimens he had seen, as a result of which Bernardi and de Lesse (1951: 141) were unable to identify syntypes.

Note 40. geyeri Aigner-Abafi, 1906: 208: status uncertain. It is not possible to synonymise this aberrant male with either M. ornata or nominotypical phoebe, since both fly in Hungary (cf. Varga 1967; Varga et al. 2005). This name was credited to Abafi-Aigner (sic) by Higgins (1941: 340).

Note 41. guevara Fruhstorfer, 1917: 19: synonym of phoebe occitanica. A lightly marked form from Spain with pale yellow ground colour, markings reduced on both wing surfaces (Higgins 1941: 340). It was synonymised by van Oorschot and Coutsis (2014: 61) with M. phoebe; its origin suggests it is synonymous with phoebe occitanica (see also Note 19).

Note 42. gurtleri Joukl, 1908: 97: status uncertain. This name was based on a single aberrant specimen, with a bright orange ground colour on the upper surface of the wings and underside wing bases that were said to be very dark. This sounds like M. ornata rather than M. phoebe, but the presence of M. ornata, although reported from Croatia (Koren and Štih 2013) has yet to be confirmed there. M. phoebe is certainly present further north than the Plitvice Lakes [TL], in Istria (Russell and Pateman 2013a, b). Overlooked by van Oorschot and Coutsis (2014).

Note 43. juliae Molina & Ocete, 1986: 869: form of Melitaea phoebe occitanica.

Note 44. koios Fruhstorfer, 1908: 194: synonym of nominotypical phoebe. Holotype examined by Bernardi and de Lesse (1951: 141) from specimens in the Museum National d’Histoire Naturelle de Paris (MNHN). Van Oorschot and Coutsis (2014: 60) gave the TL as: “Italy (S Tyrol), Switzerland (Klausen)”. Higgins (1941: 340) synonymised this large and rather dark form with nominotypical phoebe, and this was followed by van Oorschot and Coutsis (2014: 60).

Note 45. kovacsi Varga, 1967: 131: synonym of Melitaea ornata. This form has been placed in various combinations, including M. ogygia kovacsi (Varga et al. 2005) and M. ornata kovacsi (Tóth and Varga 2011). The post diapause larvae have red-brown heads (Varga et al. 2005: 67, fig. 2; Russell et al. 2007: 159, fig. 18). The present authors can find no significant morphological features to separate kovacsi from nominotypical ornata.

Note 46. leechi Rothschild, 1917: 99: Melitaea punica. A junior primary homonym of Melitaea leechi Alphéraky, 1895 (van Oorschot and Coutsis 2014: 66) (see also Note 37).

Note 47. lokris Fruhstorfer, 1908: 194: synonym of nominotypical phoebe. The upperside black pattern is more extensive than that of ottonis Fruhstorfer, 1916 (see Note 61) (Higgins 1941: 340). Type material was examined by Bernardi and de Lesse (1951: 141) from specimens in the MNHN, Paris.

Note 48. malvida Gaede, 1930, in Seitz (Supplement: 207, fig. 13d): presumed synonym of nominotypical phoebe. Gaede attributed this name to Fruhstorfer, but without a date; unable to find an original Fruhstorfer reference, Higgins (1941: 340) attributed it to Seitz (we have also failed to find any original Fruhstorfer reference, but we note that the Melitaea section of Seitz’ Palaearctic supplement was by Gaede, not Seitz). Gaede noted that malvida had pointed forewings, suggesting a form of phoebe rather than of ornata and suggested an association with form narenta (see Note 55). He also illustrated (in Seitz 1930, Supplement: plate Neptis-Argynnis, fig. d: 5) the upperside, which is not helpful for identification. Although Tóth et al. (2014: 752, fig. 1, map) indicated the presence of M. ornata in Bosnia, no locality in Bosnia was given in their specimen list (Tóth et al. 2014: 751, table 1); the present authors are not aware of any modern records of ornata from Bosnia, and a TL of Bosnia suggests synonymy with nominotypical phoebe. Thurner (1964: 34), using the name malvinda Fruhstorfer (presumably a misspelling of malvida), suggested this form was also found in the Republic of Macedonia (formerly Yugoslavia).

Note 49. mandarina Seitz, 1909: 217: synonym (provisional) of nominotypical phoebe. This very large form (Higgins 1941: 340) occurs in the eastern Palaearctic, considerably further east of the presently known eastern limit of M. ornata. Higgins (1941:340) suggested its separation from form changaica (see Note 24) was doubtful. Synonymised with phoebe by van Oorschot and Coutsis (2014: 60).

Note 50. melanina Bonaparte, 1831 (125): 159: aberration of phoebe occitanica. This male aberration had the discal ground colour of the underside of the hindwings and the submarginal lunules black. It was taken in July at Subiaco, which is only 400 m above sea level, thus it was almost certainly a specimen from a second brood, ruling out M. ornata. Not mentioned by any modern authors.

Note 51. minoa Fruhstorfer, 1917 (A. 2): 2: synonym of nominotypical phoebe. Higgins (1941: 341) treated this as a small dark race found at high levels, probably identical with nominate phoebe; van Oorschot and Coutsis (2014: 61) also placed this with M. phoebe. Type material was examined by Bernardi and de Lesse (1951: 141).

Note 52. minor Wheeler, 1903: 84: an aberration of nominotypical phoebe based on size, specimens having less than 38 mm wingspan. This was an infra-subspecific name, with no status under The Code, but for the record, the name is preoccupied by Melitaea arcesia minor Elwes, 1899 (Higgins 1941: 341). Higgins (1941: 341) suggested, and the present authors concur, that the authority was probably Wheeler as there is no reference given for Frey in Wheeler’s book.

Note 53. monilata Verity, 1919: 184: synonym of nominotypical phoebe. A large, boldly marked and bright alpine form; a specimen of this form from Simplon, Berisal, Switzerland, was figured by Higgins (1941: plate 14, fig. 1); who believed (Higgins 1941: 341) it was related to ottonis (Note 61). It is placed with nominotypical phoebe due to its TL and similarity to alternans (Note 15). Overlooked by van Oorschot and Coutsis (2014).

Note 54. monilataeformis Verity, 1919: 184: synonym of phoebe occitanica. This name was raised by Verity (1919: 184) for those specimens of tusca Verity, 1919 (see Note 96), which displayed monilata characters (see Note 53); a TL of peninsular Italy suggests synonymy with phoebe occitanica.

Note 55. narenta Fruhstorfer, 1917 (A. 2): 1, pl. 1, fig. 1: synonym of nominotypical phoebe. Fruhstorfer gave the TL as “Jablanica, Herzegovina”, which van Oorschot and Coutsis (2014: 61) wrongly interpreted as Mount Jablanica, which is on the Macedonia (FYROM)/Albania border. Seitz (1909: 207) and Higgins (1941: 341) described this as a large dark race, likening it to ottonis (see Note 61). Holotype ♂ inspected by Bernardi and de Lesse (1951: 141). Adults reared from a population of confirmed M. phoebe (i.e. final instar larvae with black heads and a white lateral stripe) from Serbia were large and dark (Peter Russell pers. obs.); it is likely that such adults are referable to narenta. Both sexes of this form were figured by Gaede (in Seitz 1930: supplement: pl. Neptis-Argynnis, figs d: 3, 4) but these were not as dark as reared specimens from Serbia. Synonymised with phoebe by van Oorschot and Coutsis (2014: 61).

Note 56. nigroalternans Verity, 1919: 184: synonym of nominotypical phoebe. An alpine form, which resembles alternans (see Note 15) but with a more extensive black pattern (Higgins 1941: 341). Overlooked by van Oorschot and Coutsis (2014).

Note 57. nigrogygia Verity, 1939: (17): synonym of phoebe occitanica. There has been some confusion related to this taxon. The TL was clearly stated by Verity (1939: (17); 1938: plate III, figs 12 and 14) to be Abbazia, Istria. At that time Istria was part of Italy but after World War 2 it became part of Croatia and the name was changed to Opatija. Higgins (1955: 118) gave the TL as “St. Dionisio, Macedonia at 800 m., gen. 2”, mistakenly using data from Verity’s postnarenta (see Note 68). Tóth and Varga (2010: 274) correctly cited the TL as ‘Opatija, Croatia’; but later wrongly as ‘Opatija, Macedonia’ (Tóth and Varga 2011: 264). Tóth and Varga (2011: 259–260), who did not examine any specimens from Croatia in their published researches on Melitaea phoebe species-group genitalia, suggested that ‘race’ nigrogygia was a subspecies of M. ornata and not of M. phoebe. Van Oorschot and Coutsis (2014: 63) also placed this taxon under ‘Melitaea telona’ (i.e. ornata) as opposed to M. phoebe, accepting the information for the TL given by Higgins (1955: 118) (John Coutsis pers. comm.). Verity (1950: 4 p.152 and Tav. 43: figs 70 and 71) figured the same two ♂♂ he figured in 1938, with the added information: ‘captured 15 May’ (year not stated) with the original locality data: ‘Abbazia, Istria’. A capture date of 15 May does not fit with second generation specimens of M. phoebe, as was suggested by Higgins (1955: 118). Russell and Pateman (2013a, b) reared a brood of M. phoebe from eggs laid by a female “nigrogygia” taken within 20 kilometres of Opatija; the larvae had black heads throughout their lives and most had an orange lateral stripe, clearly associating the taxon nigrogygia with M. phoebe occitanica, with which it is synonymised here. A study of Verity’s actual specimens may provide further enlightenment.

Note 58. nimbula Higgins, 1941: 337: synonym of phoebe occitanica. Higgins (1941: 337) raised this name for specimens of occitanica (see Note 5) with an exaggerated black pattern on the upperside. It was overlooked by van Oorschot and Coutsis (2014).

Note 59. ogygia Fruhstorfer, 1907: 310: synonym of Melitaea ornata. Recognised as a distinct species by Lafranchis (2007a, b, 2008) but considered a subspecies of M. ornata by Tshikolovets (2011), with a distribution of ‘S. and C. Greece (including Peleponnese and W. Aegean Is.); probably S.-W. Bulgaria and European Turkey’. M. ornata appears to be widespread in Bulgaria (Kolev 2015, pers. comm.). Hesselbarth et al. (1995: 1031–1033) listed over 150 locations for this species (as ‘punica telona’), all of which were in Asian Turkey. So far as the authors are aware M. ornata has not been recorded from the Greek region of Thrace, adjacent to European Turkey (Pamperis 2009: 433). The name ogygia was placed as a subspecies of M. ornata by Tshikolovets (2011: 498), as a synonym of M. punica telona (i.e. ornata) by Hesselbarth et al. (1995: 1030), and as a synonym of M. telona (i.e. ornata) by van Oorschot and Coutsis (2014: 63). Russell et al. (2007: 159, figs 16, 17) demonstrated that the larvae had red-brown heads and thus ogygia is placed as a synonym of M. ornata. The TL was given by Hesselbarth et al. (1995: 1031) as ‘Poros, Meerenge von Salamis’; the Straits of Salamis do not exist near Poros Island, nor does it feature on any of the original specimen labels (Russell and Pamperis 2011: 143). Holotype identified by Bernardi and de Lesse (1951: 140).

Note 60. ornatiformis de Sagarra, 1930: 114: synonym of phoebe occitanica. Type material taken by Querci, 24.viii.1928 at Villacabras, central Spain. Despite its nomenclatural association with ornata, geographical source clearly places this with phoebe occitanica.

Note 61. ottonis Fruhstorfer, 1917 (A. 2): 1, nota: synonym of nominotypical phoebe (a replacement name for M. phoebe var. caucasica Staudinger 1870 (see Note 22)). Higgins (1941: pl. 14, fig. 1) figured an example of this form from Simplon, Berisal, Switzerland, from which the size and the wing markings clearly suggests synonymy with nominotypical phoebe. Tshikolovets (2011: 497) treated this as a subspecies of M. phoebe, as did Tshikolovets and Nekrutenko (2012: 293) and Tshikolovets et al. (2014: 318–319). Specimens figured by Hesselbarth et al. (1995 3: Tafel 80/81: figs 30–33 ♂♂; Tafel 82/83: figs 1–4 ♀♀) from eastern Turkey, by Tshikolovets (2003: plate 24: figs 16 ♂ and 17 ♀) from Taberda, Russian Caucasus and by Tshikolovets et al. (2014: plate LX, figs 1–3 ♀♀) from Iran suggest that ottonis is best placed as a synonym of nominotypical phoebe, as van Oorschot and Coutsis (2014: 61) suggested.

Note 62. paedotrophus Bergsträsser, 1780: 14, pl. 75, figs 5–6. Synonym of nominotypical phoebe.

Note 63. parascotosia Collier, 1933: 54: Melitaea scotosia. Name based on a single ♀ specimen taken in July 1923; the author considered this subspecies to be intermediate between scotosia Butler and mandarina Staudinger. Higgins (1941: 341) considered that the name was “Probably referable to scotosia”. Lee (1982: 46) placed scotosia Butler [TL: Tokyo, Japan] as a subspecies of M. phoebe. However, Tuzov et al. (2000: 2: 74), Gorbunov and Kosterin (2007: II: 85) and van Oorschot and Coutsis (2014: 67) synonymised the name with Melitaea scotosia Butler, which occurs in the eastern Palaearctic. Although originally described as a subspecies of M. phoebe, it does not appear to be associated with any of the three taxa (phoebe, ornata, punica) dealt with in this paper.

Note 64. parva Gerhard, 1882: 126: synonym of nominotypical phoebe. A bright “second generation” form, reared from a larva – colour and host-plant unknown. Higgins (1941) and van Oorschot and Coutsis (2014) overlooked this form.

Note 65. parva Caradja, 1895: 47: probable synonym of nominotypical phoebe. A small, brightly marked variety of the first generation (Higgins 1941: 341). A larva from Transylvania, Romania, having typical characters (black head with white lateral stripe) of nominotypical phoebe was figured by Russell et al. (2007: 159, fig. 14). Székely (2008: 175–176) included reports (unconfirmed by larval head colour) by T. Hácz of M. punica telona (= ornata) from Transylvania and North-Dobrudja in Romania; however, these records were reported later by Hácz (2012: 73) as M. phoebe. Not mentioned by van Oorschot and Coutsis (2014). Since both this and the previous entry are infrasubspecific, they are not covered by The Code.

Note 66. pauper Verity, 1919:183: synonym of phoebe occitanica. Described as a small, lightly marked form with pale ground colour, the usual summer brood form of tusca Verity, 1919 (Higgins 1941: 341 and pl. 14, fig. 3) (see also Note 96).

Note 67. phoebina Turati, 1919: 222: synonym of Melitaea ornata. A small mountain form (Aspromonte, above 1400 m) rather dark and heavily marked, related to totila Stauder, 1914 (Higgins 1941: 341) (see Note 92). According to Turati (1919: 222) there is no second generation of this form, which he considered similar to that from Ficuzza, Palermo, Sicily (see Note 32). The TL is outside the range of M. phoebe, which has not been observed south of Monte Martinellal, Cosenza, Calabria, at the much lower elevation of 880 m (cf. discussion on altitudinal separation in Italy in Russell and Pateman 2011: 28) from where 5♂♂ were taken by the first author (identification confirmed from genitalia, club shaped antenna and underside hindwing pattern). Overlooked by other authors, including van Oorschot and Coutsis (2014).

Note 68. postnarenta Verity, 1939: (17): synonym of nominotypical phoebe. Verity (1939: (17) gave this name to small second generation specimens of M. phoebe, resembling emipauper (see Note 31). The TL and details of collection for this form were mistakenly attributed by Higgins (1955: 118) to nigrogygia (see Note 57). Resemblance to emipauper is superficial. Overlooked by van Oorschot and Coutsis (2014).

Note 69. postogygia Verity, 1939: (16): synonym of nominotypical phoebe. Verity (1939: [16]) gave this name to a small form flying in the hills above Thessalonica in August; close association with the name ogygia (i.e. M. ornata – see Note 64) is misleading. Higgins (1955: 118) included the name in his list of synonyms of M. phoebe and indicated a similarity with parva (see Notes 64 and 65) and pauper (see Note 66). A second generation form (M. ornata is single-brooded – see Note 2) from central Greece places this taxon with nominotypical phoebe. It was overlooked by van Oorschot and Coutsis (2014).

Note 70. postvirgilia Verity, 1950: 154: synonym of nominotypical phoebe. The second generation of the Alpine first generation form virgilia (see Note 100). Not listed by Higgins (1941, 1955) or any recent authors.

Note 71. pseudosibina Alberti, 1969: 192, Taf. 1, figs 1c and 2c.: synonym of nominotypical phoebe. Synonymised with nominotypical phoebe by Hesselbarth et al. (1995: 1028), and with “M. phoebe” by van Oorschot and Coutsis (2014: 61). Judging from the paratypes figured by Alberti (1969: Taf. 1, figs 1c and 2c) and the specimen figured by van Oorschot and Coutsis (2014: plate 12, fig. 24), which has clubbed antenna and hindwing underside arcuate submarginal markings reaching the intervening veins, this is correct. Tshikolovets (2011: 497) placed it as a synonym of Melitaea phoebe ottonis (see Note 61), as did Tschikolovets and Nekrutenko (2012: 293).

Note 72. punicapowelli Oberthür, 1915: fig. 2338: synonym of Melitaea punica. Specimens of M. punica which have the black pattern partly obsolete (Higgins 1941: 342).

Note 73. punicata Ragusa, 1919: 150: synonym of Melitaea ornata. Equated to emipunica (see Note 32) by Higgins (1941: 342).

Note 74. reliquiae Korb et al., 2015: 143 and plate VI: synonym of Melitaea ornata. Information on the populations in the Volgograd region was first published by Kuznetsov and Stradomsky (2010) under the name Melitaea telona and later by Russell and Kuznetsov (2012) under the name M. ornata.

Note 75. rostagnoi Turati, 1920: 223: synonym of phoebe occitanica. A small second generation form, probably much the same as emipauper Verity (see Note 31) and autumnalis Fruhstorfer (see Note 17) (Higgins 1941: 342). Synonymised with M. phoebe by van Oorschot and Coutsis (2014: 61).

Note 76. rovia Fruhstorfer, 1919: 169: synonym of nominotypical phoebe. According to Higgins (1941: 342) this is a low elevation form with reduced black markings. A holotype and allotype were examined by Bernardi and de Lesse (1951: 141). Synonymised with M. phoebe by van Oorschot and Coutsis (2014: 61).

Note 77. rubialesi Gómez Bustillo, 1973: 36: form of Melitaea phoebe occitanica.

Note 78. rubrofasciata Gušić, 1922: 95: probably a synonym of nominotypical phoebe: Higgins (1941: 342) Noted this as a small form with a deficiency of black pattern on the discal area of the wings. However, the name seems to indicate the presence of some red colour on the wings. Although Koren and Štih (2013) recorded M. ornata from five localities in Croatia, one of which was near Zagreb, the identity of the species has been questioned (Koren pers. comm.) The first author visited two of the locations concerned in May 2015 and considered that the biotope was unsuited to M. ornata. Podsused (the TL) is on the banks of the River Sava at c. 125 m above sea level and appeared on recent inspection to be encompassed by industrial buildings (Russell pers. obs.); it would seem unlikely that either species would be extant currently in that locality.

Note 79. sarvistana Wiltshire, 1941: 473, fig. 3: Melitaea sarvistana. Originally described as a race of M. phoebe based on two male specimens; a large form, with black submarginal lunules complete on both wings, other markings faint with nearly obsolete discal markings; on the underside of hindwings the black markings are prominent (Wiltshire 1941). Wiltshire (1946: 25, plate 1: figs 1 and 2)), from an examination of the genitalia, elevated this to species status. Higgins (1955: 117, pl. I, fig. 17 pl. II, fig. 17) also considered it a distinct species. Eckweiler and Hofman (1980: 10), Racheli (1980: 80–81), Koçak et al. (1997: 4), Nazari (2003), Kolesnichenko (2007: 30), van Oorschot and Coutsis (2014: 69 and pl. 14, figs 20–22) and Tshikolovets et al. (2014: 321 and pl. LX, figs 13–15, 18) all followed Wiltshire in recognising sarvistana as a distinct species. The present authors have no personal experience of this taxon but it appears to be different from any examples of the taxa under consideration; its inclusion here is only because it was originally described in association with M. phoebe.

Note 80. saturata Staudinger, 1892: 323: synonym of nominotypical phoebe. A large brightly coloured form resembling many mountain forms of phoebe (Higgins 1941: 342). It occurs in the eastern Palaearctic and, since the presently known eastern limit of the distribution of M. ornata is Kazakhstan, southeast of the Ural Mountains, placement with nominotypical phoebe seems appropriate. Synonymised with M. phoebe by van Oorschot and Coutsis (2014: 60). Korb (2011: 158) identified Melitaea specimens from N Tian-Shan as M. phoebe saturata. The TL of saturata is Mongolia, some 2500 km northeast from Tian-Shan. Korb et al. (2015: 142–143, Col. pl. VI, figs 3 and 4), who then considered that M. phoebe was absent from Tian-Shan, reassessed this population as M. ornata; we consider saturata a synonym of M. phoebe phoebe.

Note 81. seminigra Delahaye, 1909: 10: aberration of phoebe occitanica. This aberrant female specimen, with almost black forewing uppersides, was taken in June at Pignerolles, Maine et Loire in west central France and thus outside the ranges of both nominotypical phoebe and M. ornata. Higgins (1941: 342) stated that he did not view the original publication and thus could make no comment on this name. It has not been mentioned by any recent author.

Note 82. sextilis Jachontov, 1909: 285: synonym of nominotypical phoebe. A small second generation form of caucasica (see Notes 22, 61) taken in the southern Caucasus in August. Higgins (1941: 342) and van Oorschot and Coutsis (2014: 60) placed this with M. phoebe.

Note 83. sibina Alphéraky, 1881: 400, Tabl. XIV fig.13: status unclear (distinct species/synonym of nominotypical phoebe). This taxon is distributed mainly outside the western Palaearctic, with a western distribution limit in the Republic of Kazakhstan (Tshikolovets 2003: 328). Originally described as a variety of M. phoebe, it was given species status by Higgins (1941: 349, plate 15: figs 5, 6, 11 and 12) and this has been followed by some recent authors (for example: Tshikolovets 2003: 328–329, 2005: 338; van Oorschot and Coutsis 2014: 65–66). It does not appear to be directly associated with any of the three taxa dealt with in this paper. We note that Tóth and Varga (2011) and Tóth et al. (2014) were unable to separate it from M. phoebe using molecular or morphometric procedures.

Note 84. sterlineata Turati, 1920: 223, Tav. II, figs 10–12: synonym of phoebe occitanica. Although placed by Turati as an aberration of phoebina (= ornata, see Note 64), the specimens were taken by GC Krüger, at 800 m altitude, in September 1909; it must therefore represent a second or even third generation form, which precludes it from being ornata, which is univoltine.

Note 85. streltzovi Kolesnichenko & Yakovlev, 2004: 103: synonym of nominotypical phoebe. Distributed along the south-western slopes of the Mongolian Altai. All specimens taken in the first part of July, flying in mesophilous grasslands and river valleys. The figures (Kolesnichenko and Yakovlev 2004: figs 10, 11 on plates V and VI) show both sexes are heavily marked with a pale background on the upper surfaces of both fore- and hindwings. The club shaped antenna and the arcuate submarginal markings appearing to reach the intervening veins (see Table 1), suggest association with M. phoebe rather than M. ornata. Synonymised with M. phoebe using van Oorschot and Coutsis (2014: 61)

Note 86. subcorythallia Verity, 1928: 162: synonym of phoebe occitanica. “The second generation of France” (Higgins 1941: 342).

Note 87. suboccitanica Verity, 1928: 162: synonym of phoebe occitanica. “The first generation of France” (Higgins 1941: 342).

Note 88. subtusca Verity, 1952: 349: synonym of phoebe occitanica. Verity (1952: 349) referred this form from southeast France to tusca from central Italy (see Note 96), and it is placed with phoebe occitanica as a result. M. ornata from Var, France occurs in a very different phenotype from the form of phoebe occitanica occurring in central Italy (Verity 1951: plate 44, figs 1–16), the former being much darker in colour and having triangular submarginal lunules (Russell et al. 2007: 162 fig. 52). Overlooked by van Oorschot and Coutsis (2014).

Note 89. sylleion Fruhstorfer, 1917 (A. 2): 2: synonym of nominotypical phoebe. Higgins (1941: 342) considered this form to be inconsistent. The holotype and allotype were inspected by Bernardi and de Lesse (1951: 141). It was placed by van Oorschot and Coutsis (2014: 61) as a synonym of M. phoebe.

Note 90. tatara Krulikovsky, 1891: 236: status uncertain (possibly a hybrid). Spelt tartara (sic) by Higgins (1941: 342) but subsequently corrected (Higgins 1944: 46). The origin of this name refers to Tatastan, a Russian Province in which Casan, the TL, is located. Although Krulikovsky placed it under M. phoebe, M. ornata was not at that time established as a species. Higgins (1941: 342) said: ‘An example in which there is a double black line across both wings parallel to the outer margin’ but this is not helpful to place it with either species. Krulikovsky’s figure (1890: 236, VIII, fig. g) does not allow identification; in fact Krulikovsky himself suspected that it was a hybrid between M. phoebe and M. athalia. Having later observed a ♂ M. phoebe coupling with a ♀ M. arduinna (Esper, [1783]), Krulikovsky (1897: 321), restated his suspicion that tatara was a hybrid. It is noted that hybrids have been recorded between M. ornata and M. phoebe by Bálint and Ilonczai (2001: 217) in Hungary and by Russell et al. (2014: 140, figs 7–9) in Slovenia; since both species probably occur in this area to the west of the Urals in the Russian Federation, a hybrid origin remains a possibility.

Note 91. telona Fruhstorfer, 1907: 310: synonym of Melitaea ornata (but see below). The holotype and allotype were examined by Bernardi and de Lesse (1951: 140). This name was placed as a subspecies of M. ornata by Tshikolovets (2011: 499) and by Tshikolovets and Nekrutenko (2012: 295). This is the name used by a number of authors for what is now known to be M. ornata, including the first author (Russell 2008; Russell and Pateman 2011), prior to our present understanding of the range of M. ornata, which led to the recognition that ornata and telona were conspecific. Russell et al. (2007: 159, fig. 15) demonstrated that the larva of telona from its TL has a red-brown head; larvae of ornata from Volgograd region, Russia, are similarly coloured and also has a red-brown head (Russell and Kuznetsov 2012: figs 1–3), suggesting synonymy with M. ornata. However, recent molecular analysis by Tóth et al. (2014) apparently suggests that telona may represent a species distinct from ornata; only two samples of telona from Lebanon, the origin of the ‘voucher specimen’ used as an example of telona by Wahlberg and Zimmermann (2000) for their mtDNA sequencing, were included in their analysis. Until this is resolved, it is considered prudent to retain telona as a synonym of M. ornata. Rather confusingly, van Oorschot and Coutsis (2014: 63) considered telona a distinct species and placed ornata as a synonym of M. phoebe.

Note 92. totila Stauder, 1914: 373: synonym (provisional) of Melitaea ornata. The first author visited Monte Cocuzzo, the TL, on a number of occasions but, in spite of the presence of a known host-plant (Centaurea deusta Ten.: Russell and Pateman 2011) only discovered one worn ♀ at ca. 1200 m, which unfortunately died prior to ovipositing. It appeared from its hindwing markings and spatulate antenna to be M. ornata. Also, a single ♂ was taken on Monte Mancuso, Calabria, some 24 km to the south, which from an examination of genitalia and external morphology, was almost certainly M. ornata. This form is therefore provisionally placed with M. ornata.

Note 93. tremulae Piller & Mitterpacher, 1783: 69, Taf. 4: figs 1 and 2: synonym (provisional) of nominotypical phoebe. The TL of Croatia, from where there have been no substantiated reports of M. ornata, strongly suggests association with phoebe phoebe. Hesselbarth et al. (1995: 1028) synonymised this name with M. phoebe phoebe. Not mentioned by any more recent authors.

Note 94. tungana Seitz, 1909: 216: synonym (provisional) of nominotypical phoebe. The specimens were described by Seitz (1909: 216) as very melanic but the specimens examined by Higgins (1941: 342) showed that this character was variable in the Sayan Mountains (the TL) and suggested that many of these specimens were close to monilata Verity (see Note 53) and other Alpine forms (see Higgins 1941: 334). The form tungana has a distribution in the eastern Palaearctic and outside the presently recorded distribution of M. ornata, the authors provisionally place tungana with M. phoebe. Overlooked by van Oorschot and Coutsis (2014).

Note 95. tungusa Herz, 1899: 240: synonym (provisional) of nominotypical phoebe. A small form with obscure markings, in appearance somewhere between var. caucasica Staudinger (see Note 22) and M. ornata (see Note 2). Synonymised with M. phoebe by van Oorschot and Coutsis (2014: 60). Since it occurs in the eastern Palaearctic, it is synonymised with nominotypical phoebe until further information becomes available.

Note 96. tusca Verity, 1919: 183: synonym of phoebe occitanica. Described by Verity (1909: 183) as a form with bright orange upperside ground colour and reduced black markings. Higgins (1941: 342) considered this to be a first (spring) brood form from central Italy, and was of the opinion that the names emipauper Verity, 1919, pauper Verity, 1919 and probably autumnalis Fruhstorfer, 1919 referred to the second or third (summer) broods of tusca (see Notes 31, 65 and 17, respectively). Placed here as a synonym of phoebe occitanica largely due to its geographical location in peninsular Italy. This name was overlooked by van Oorschot and Coutsis (2014).

Note 97. uclensis Melcón, 1910: 219: aberration of phoebe occitanica. Described as an aberration of occitanica with the upperside black marginal semi-lunules separated from the black marginal line by red ground colour. Its origin in central Spain clearly places it with occitanica. Overlooked by van Oorschot and Coutsis (2014).

Note 98. ufensis Krulikovsky, 1902: 555 (footnote): synonym of Melitaea ornata. A replacement name for uralensis Krulikovsky, 1897 (see Note 99); van Oorschot and Coutsis (2014: 60) incorrectly placed this as a synonym of M. phoebe.

Note 99. uralensis Krulikovsky, 1897: 3: name preoccupied by Melitaea arduinna uralensis Eversmann, 1844. Replaced with ufensis by Krulikovsky (1902: 555 footnote). Van Oorschot and Coutsis (2014: 60) incorrectly placed this as a synonym of M. phoebe (see Note 98).

Note 100. virgilia Fruhstorfer, 1917 (A. 2): 2: synonym of nominotypical phoebe. Higgins (1941: 343) treated this as a large race with pale ground colour and black markings reduced, although he recognised that these features were not constant. The relatively larger than average size and its TL in the French Alps places this taxon with nominate phoebe, with which it was placed by van Oorschot and Coutsis (2014: 61). Holotype and 5 ♀♀ paratypes were examined by Bernardi and de Lesse (1951: 141).

Note 101. wagneri Wnukowsky, 1929: 222: replacement name for alatauica Wagner, 1913 (see Note 11).

Note 102 yagii Nire, 1917: 146, including fig. 2: Melitaea scotosia.This taxon is confined to the eastern Palaearctic. The name yagei (sic) was synonymized with M. scotosia Butler, 1878 by Higgins (1941: 343). This synonymy and misspelling were followed by van Oorschot and Coutsis (2014: 67). Although originally placed with M. phoebe, it does not appear to be associated with any of the three taxa (phoebe, ornata, punica) dealt with in this paper.

Note 103. zagrosi Tóth & Varga, 2011: 265: synonym (provisional) of Melitaea ornata. This name was raised as a distinct species based on male and female genitalia and underside wing markings. However, it would appear from Tshikolovets et al. (2014: 320, map) that the type locality of this form is within the distributional area of abbas, which they elevated to a subspecies of Melitaea ornata. Van Oorschot and Coutsis (2014: 64) discussed the status of zagrosi at some length, referring to the unreliability of wing markings, which has been demonstrated in Melitaea taxa by Jugovic and Koren (2014), and genitalia preparations when placed in covered slides creating distortion. They concluded that there was insufficient evidence to support the erection of a new taxon and classed it as ‘status incertus’. Tshikolovets et al. (2014: 320) synonymised it with M. ornata abbas (i.e. ornata) (see Note 7). The elevations at which these two forms occur (zagrosi, 300 m; abbas, 1500–2500 m) may be significant. Until additional evidence becomes available, it is provisionally placed with M. ornata.