Parnassiinae Duponchel, [1835]

Introduction

The subfamily Parnassiinae is a group of essentially Palaearctic butterflies that live in a variety of habitats, ranging from arid deserts (Hypermnestra) to humid forests (Luehdorfia), lowland meadows (Zerynthia), and high alpine habitats (Parnassius). A few species of the genus Parnassius constitute the only representatives of the subfamily in the Western Hemisphere, where they occur in the Nearctic region (Opler and Warren, 2003).

These butterflies have been studied since the time of Linnaeus (1758), who named, among others, the magnificent Parnassius apollo. Interest in the group grew in the 18th and early 19th century when more material became available from the Far East and the Himalayas (Smart, 1976). The breathtaking beauty of Bhutanitis species and the variability of the genus Parnassius quickly made them popular among butterfly collectors who bragged about rare and unusual specimens in their collections (Salmon, 2000). At some point in the nineteenth century, these butterflies gained so much popularity that many European museums deployed expeditions to the Himalayan region solely in search of rare and undiscovered species of Parnassiinae (Talbot, 1939). Of the three parties of collectors sent in search of Bhutanitis lidderdali between 1868 and 1890 by the British Museum, the first was plundered by natives, the second was stricken by fever and one of its members died, and the third had a man killed by a tiger. All three returned without success, although further specimens of this species became available a few years later in the 1890s (Talbot, 1939).

The subfamily includes eight extant genera that, based on most recent molecular studies, can be grouped in three tribes: Parnassiini (Hypermnestra, Parnassius), Zerynthiini (Sericinus, Bhutanitis, Zerynthia, Allancastria), and Luehdorfiini (Luehdorfia, Archon) (sensu Nazari et al., 2007; also see Stekolnikov and Kuznetsov, 2003; Omoto et al., 2004; Katoh et al., 2005). Parnassius has the highest number of species among the genera in the subfamily, and depending on the checklist, between 38 to 47 species are recognized, each with many subspecies and individual forms (Bryk, 1935; Collins and Morris, 1985; Weiss, 1991-2005; Häuser et al., 2005). The systematic position of two fossil taxa, Thaitites ruminiana and Doritites bosniackii, has been difficult to resolve partly due to incomplete character preservation (Hancock, 1983, Nazari et al., 2007).

Characteristics

Putative synapomorphies for Parnassiinae include (Ehrlich, 1968; Miller, 1987; Nazari et al., 2007):

• Asymmetrical pretarsi,
• Narrow and heavily sclerotized aedeagus,
• Heavily sclerotized ostial region in females,
• Elongate third segment of labial palpus,
• Incurved middle discocellular vein on the forewing.

Putative synapomorphies for Parnassiini (Hypermnestra , Parnassius) are (Hancock, 1983; Miller, 1987; Nazari et al., 2007):

• Solitary larvae,
• Cocooned pupae,
• Scaled antennae,
• Sclerotized patagia,
• No mid-tibial spurs on hind legs,
• Simple (unforked) precostal vein on the hindwing.
  

Hypermnestra possesses many autapomorphies that suggest a deep divergence with Parnassius (Nazari et al., 2007).

Putative synapomorphies for Zerynthiini (Sericinus , Bhutanitis , Zerynthia , Allancastria) include (Hancock 1983; Miller 1987; Nazari et al., 2007):

• Larvae equipped with fleshy segmental tubercles with setae,
• Slender pupae,
• No scales on tibia and tarsi in adults.

The tribe Luehdorfiini (sensu Nazari et al., 2007) was recently revised based on DNA evidence from five mitochondrial and two nuclear genes, and there are no morphological characters known to unite Archon and Luehdorfia (Nazari et al., 2007).

Discussion of Phylogenetic Relationships

Three tribes are recognized within Parnassiinae: Parnassiini (including Hypermnestra and Parnassius), Zerynthiini (including Allancastria, Sericinus, Zerynthia, and Bhutanitis), and Luehdorfiini (including Archon and Luehdorfia) (sensu Nazari et al., 2007). Most previous morphological classifications, however, placed Archon with Parnassiini and Luehdorfia with Zerynthiini, recognizing only two tribes (Ehrlich, 1958; Munroe, 1961; Ackery, 1975; Hancock, 1983; Igarashi, 1984; Collins and Morris, 1985; Häuser et al., 2005). Some of these studies have considered these two tribes to be separate subfamilies (Bryk, 1934; Talbot, 1939; Ford, 1944a, b; Eisner, 1974; Higgins, 1975; Chunsheng, 2001) or even families (Clench, 1955; Hemming, 1960; Eisner, 1974). although most recent authors discount this idea (e.g. Häuser et al., 2005).

The correct positions of the genera Archon, Hypermnestra and Luehdorfia have been disputed; Archon has sometimes been included in Zerynthiini (Eisner, 1974; Higgins, 1975) or in a separate sub-tribe within Parnassiini (Koçak, 1989). Häuser (1993) suggested a separate subfamily for Hypermnestra based on a number of morphological and ecological autapomorphies, a view previously expressed by Dujardin (1965), Hiura (1980), and Korshunov (1990) (as reported by Korb, 1997). A recent study of genitalic characters (Stekolnikov and Kuznetsov, 2003) recognized the tribe “Hypermnestriini Hiura 1980” and gave subfamily status (Luehdorfiinae Tutt, 1896) to Luehdorfia based on putatively primitive genitalic characters (also see Ford, 1944b).

The molecular phylogeny of the subfamily has been unresolved for a long time, due in part to incomplete sampling of previous molecular studies (Caterino et al., 2001; Omoto et al., 2004; Katoh et al., 2005). However, a recent study (Nazari et al., 2007) using evidence from five mitochondrial and two nuclear genes in conjunction with 236 morphological characters, has determined three tribes within the subfamily Parnassiinae, as discussed above.

The systematic position of two fossil taxa, Thaitites ruminiana and Doritites bosniackii, has remained largely unresolved (Hancock, 1983), although a close affinity between Doritites and Archon has recently been found (Nazari et al., 2007).

References

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