Pseudoneorina coulleti

Jong, Rienk De, 2017, Fossil butterflies, calibration points and the molecular clock (Lepidoptera: Papilionoidea), Zootaxa 4270 (1), pp. 1-63 : 24

coulleti . Pseudoneorina coulleti Nel & Descimon, 1984

Fig. 6.

Nymphalidae : Satyrinae (subdivisions not yet generally agreed upon).

France, Dép. Alpes-de-Haute-Provence, Céreste and Dauphin; Rupelian (= "Stampien"), early Oligocene.

Depositories: holotype from Céreste in Collection Nel, La Ciotat, Bouches-du-Rhône, France (nr 2486) ; paratype from Céreste in Collection Coullet, Barrème, Alpes-de-Haute-Provence, France ; additional specimen from Dauphin in Collection Michel Henrotay (Liège, Belgium).

Published figures: Henrotay (1986); Murata (1998: Figs 7, 8); Nel & Descimon (1984: Figs 1–5 ); Nel & Nel (1986: Fig. 2 e).

Three specimens, in varying states of preservation. Eyes naked. Forewing without vein 2A; radial formula 1, 2, 3+(4+5); stalk of R3-R5 and M1 originating from same spot at upper corner of cell; M2 at its base much closer to M1 than to M3; cell closed; base of Cu lightly swollen; rounded spot between M1 and M2; darker area between base, cubitus, M3 and hind margin, extending from base to about two-thirds towards termen; outer margin probably straight. Hindwing with Sc+R1 arising far from base: distance from base to Sc 3.6 mm, from Sc to Rs 4.5 mm; cell closed; round dark spot between Cu1 and Cu2 (according to the original description and in their photograph Figure 2 A and in the reconstruction in Figure 5 ; in the description, it is given as between Cu1 and M3); outer margin possibly extended tail-like at the end of M3 (as it is in extant Neorina ; in fig. 5 at the end of Cu1), but since the relevant part of the wing is absent in the fossil, this is speculative.

In their well-documented discussion the authors conclude that the fossil is close to the recent genus Neorina ( Nymphalidae , Satyrinae ), but differs in the origin of Sc in the hindwing being further from base than in any recent satyrine butterfly: in Pseudoneorina Sc originates at 44.4% of distance from base to Rs, further than in any extant species, in Neorina it originates at almost 22% of this distance. Apparently this feature was an autapomorphic character that was lost with the extinction of the taxon.

The venational characters of the fossil as well as the position of the rounded spots (in extant satyrines eyespots), although agreeing with extant Neorina , can be found in other extant genera as well, e.g. Zophoessa (considered a subjective synonym of Lethe [subtribe Lethina of Satyrini ] by Wahlberg & Peña 2015), but to belong to another tribe, Elymniini , in the classification accepted at the Natural History Museum, London, www.nhm.ac.uk/ our-science/data/butmoth/, which follows the database Lamas_Genera_ 04ii08. xls. Wahlberg & Peña (2015) classify Pseudoneorina within the subtribe Lethina , and Neorina as belonging to another tribe, Zetherini . Earlier Neorina was also placed outside Satyrini , as sister taxon to Penthema , in a clade that is sister taxon to Amathusiini ( Wahlberg et al. 2009). According to Miller (1968) Neorina belongs to the Lethini, in which it, with its closest relatives, is mainly diagnosed by possessing naked eyes, a character found frequently throughout the butterflies and not very indicative of relationship. As long as there are no apomorphies known that unambiguously support relationships, I would hesitate to classify the fossil either as belonging to Satyrini or to Zetherini (see also Figures 18–20 under Neorinopsis sepulta , for the still unsettled interrelationhips among the tribes of Satyrinae ).

A fossil from another locality about 18 km to the east, Dauphin (Dép. Alpes-de-Haute-Provence), Lower Oligocene, Stampien (= Rupelian), ca. 30 Ma, was figured but not named by Henrotay (1986). It was recognized as belonging to the same taxon by André Nel, co-author of the name (pers. comm. Henrotay, 16 Dec. 2015).

Figures

FIGURE 1. Localities of fossil butterflies plotted ion a modern map. See Table 1 for further, complementary information.

FIGURE 2. Apomorphies, their placement on a tree, and the assignment of calibration points. See text for explanation.

FIGURES 3 – 4. Wing venation of (A) Pareronia Bingham, 1907 (Pieridae), and (B) Hasora Moore, 1881 (Hesperiidae). See text for explanation.

FIGURE 5. Ƒanessa amerindica Miller & Brown, 1989, holotype, collections of Invertebrate Paleontology, University of Florida. Photo by Roger Portell. Fig. 6. Pseudoneorina coulleti Nel & Descimon, 1984. France, Dép. Alpes-de-Haute- Provence, Dauphin, specimen in coll. M. Henrotay; photo by courtesy of M. Henrotay. Fig. 7. Archaeolycorea ferreirai Martins-Neto, 1989. Fig. 4 A of original description. Fig. 8. Oligodonta florissantensis Brown, 1976, holotype, collections of Invertebrate Paleontology, University of Florida. Photo by courtesy of Akito Kawahara. Fig. 9. Neorinella garciae Martins- Neto, Kucera-Santos, de Moraes Vieira & de Campos Fragoso, 1993. Fig. 3 A of original description. Fig. 10. Protocoeliades kristenseni de Jong, 2016, holotype, Zoologisk Museum, Copenhagen, Denmark. Photo by Department of Paleobiology, National Museum of Natural History, Washington DC, USA.