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FIGURE 24 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 24. Hybrid #15 (dorsal, ventral). Peru: Loreto, Yurimaguas, collected in July, 1919 by O. Michael. Currently housed in the Holzinger collection in NMW. This specimen is the holotype of H. elevatus f. nöldneri Neustetter, 1938. Mallet et al. (2007) interpreted this as a F1 H. elevatus pseudocupidineus (Fig. 3) x H. hecale versicolor Weymer 1894 (a synonym of H. hecale felix, Fig. 25), backcross, presumably based on the orange patches at the base of the FW yellow band and at the outer margin of the FW.

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FIGURE 23. H. elevatus roraima Turner, 1966 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 23. H. elevatus roraima Turner, 1966 (dorsal, ventral). Venezuela: Bolivar, Via Roraima km 7. (image source: https://cliniquevetodax.com/Heliconius/pages/elevatus%20roraima).

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FIGURE 12 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 12. Hybrid #10 (dorsal; ventral). Brazil: Para, Obidos, 1985. Neukirchen collection (FLMNH). Interpreted by Mallet et al. (2007) to be a possible F1 hybrid of H. numata superioris Butler, 1875 (Fig. 5) and H. melpomene nr. meriana (Fig. 9); also suggested to be similar to hybrid #12 (Fig. 15, see below). Presumably the "hybrid" features exhibited by this specimen are the transverse orange bar and the v-shaped marks beneath it on the HW. However, the range of H. melpomene meriana does not extend to the Amazon (Rosser et al. 2012). It is replaced there by H. melpomene melpomene (Fig. 6), which has a red FW band and no markings on the HW. Thus, the hypothesized cross is biogeographically implausible. Furthermore, there are as many as five different forms of H. numata co-occurring in the lower Amazon that exhibit most of the features of this specimen (see Hybrid #8). As noted in the introduction, the Neukirchen collection is somewhat problematical as a source of reliable data, as many of its unusual specimens have dubious provenances and were likely purchased from commercial dealers.

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FIGURE 22. H. elevatus zoelleri Neukirchen, 1990 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 22. H. elevatus zoelleri Neukirchen, 1990 (dorsal). Venezuela: Amazonas, Puerto Ayacucho. R. Mattei coll. (image source: https://cliniquevetodax.com/Heliconius/pages/elevatus%20zoelleri). Note the scalloped edges of the wings.

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FIGURE 16. H. hecale vetustus Butler, 1873 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 16. H. hecale vetustus Butler, 1873 (dorsal, ventral). French Guiana: Montagne de Kaw Pk 37.5, leg. Y. Lever. (image source: https://cliniquevetodax.com/Heliconius/pages/hecale%20vetustus.html)

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FIGURE 1 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 1. Heliconius wing regions (left) and venation (right) discussed in the specimen accounts below. Abbreviations: FW = forewing; HW = hindwing; Hu = humeral vein; Sc = subcostal vein; R1–R5 = radial veins 1–5; RS = radial sector vein; M1–M3 = medial veins 1–3; Cu1, Cu2 = cubital veins 1 and 2; A1+2, A3 = anal veins 1–3. Wing outline modified from Papa et al. (2013, fig. 2).

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FIGURE 7. H. numata mavors Weymer, 1894 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 7. H. numata mavors Weymer, 1894 (dorsal). (image source: https://cliniquevetodax.com/Heliconius/pages/ numata%20mavors.html

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FIGURE 4 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 4. Hybrid #8 (dorsal, ventral). Brazil: Rio Arapiuns [near Santarém], 1980, leg. H. Holzinger[?], Holzinger collection, NMW. Discussed and illustrated in (Holzinger & Holzinger 1994, p. 94; plate 24 fig. 5c). Mallet et al. (2007) interpreted this specimen as an H. numata superioris (Fig. 5) x H melpomene melpomene (Fig. 6) F1, based on the subapical "red bar" on the forewing. The color of this element appears quite similar, at least in these photographs, to other orange areas of the fore- and hindwings. There are described forms of H. numata from Peru with subapical orange markings distal to a yellow band (H. staudingeri Weymer, 1894; H. aristiona staudingeri f. lutea Neustetter, 1931) considered intraspecific hybrid forms of H. numata by Lamas & Jiggins (2017). H. numata mavors (Fig. 7), which has no yellow bands or spots, occurs in the lower Amazon, along with several other polymorphic forms of H. numata with different wing patterns. This specimen could represent an unusual recombinant phenotype of supergene alleles that occur within H. numata (Joron et al. 2011), or even an H. numata x H. pardalinus hybrid, which might offer an explanation for the orange chevrons on the HW. Finally, in an obituary for Ruth Holzinger, Aspöck (1996) reported that neither she nor Helmuth Holzinger ever ventured outside of Europe. Thus, all of the Heliconius specimens in their collection (donated to the Naturhistorisches Museum in Vienna; Lödl et al. 1997) were obtained by gift, trade, or purchase from commercial dealers such as Hermann Gerstner.

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FIGURE 15 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 15. Hybrid #12 (dorsal; ventral). Guyane: St. Laurent. Holotype of Heliconius seraphini Talbot, 1932, from the famous hybrid zone in St. Laurent, French Guiana, collected prior to 1932 (specimen in NHMUK). Interpreted by Ackery and Smiles (1976) as H. melpomene thelxiope (Hübner, [1806]) x H. numata numata, and interpreted by Mallet et al. (2007) to be an F1 hybrid between H. hecale vetustus (Fig. 16) and H. melpomene interspecific hybrid (H. melpomene melpomene (Fig. 6) x H. melpomene thelxiopeia (Fig. 17). The locality data suggest that Mallet et al.'s parental forms should be preferred. Other than the (probably postmortem) discoloration of the yellow band, the forewing is quite hecale-like. The pointed distal ends of the FW yellow rays and the with of the HW. orange rays are reminiscent of H. melpomene thelxiopeia. However, the concave proximal edges of the (formerly) yellow FW spots and the presence of rays and submarginal white spots on HWV are more reminiscent of H. elevatus (the locally sympatric race is H. elevatus bari, Fig. 18) than of H. melpomene thelxiopeia. Given the equivocal phenotype, it is more parsimonious to infer a H. hecale x H. elevatus cross on phylogenetic grounds.

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FIGURE 11. H. numata aurora Bates, 1862 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 11. H. numata aurora Bates, 1862, "typical" form (dorsal, ventral). Peru: Loreto, Iquitos. (image source: https:/ /cliniquevetodax.com/Heliconius/pages/numata%20aurora.html). Other variants of this race are more melanic, lacking the yellow apical band on the FW.

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FIGURE 21 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 21. Hybrid #14 (dorsal, ventral). Colombia: Guainía, Puerto Inírida, 03°52' 16" N, 67°53'31" W, 100m. 14 Jan. 1974, leg. E. Schmidt-Mumm (IAvH). Mallet et al. (2007) interpreted this as a potential H. elevatus zoelleri (fig. 23) x H. hecale backcross to H. elevatus, based on the scalloped edge of the hindwing (they suggested that alternatively, it could be a variant of H. elevatus, but nevertheless counted it as an interspecific hybrid in their tally). According to Rosser et al. (2012), the only other silvaniform occurring at the collection locality is H. numata aurora (Fig. 11). This specimen is likely to be just a variant of H. elevatus, several other forms of which have yellow subapical spots on the forewing (cf. H. elevatus bari, Fig. 18), or the forewing red dennis interrupted by a silvaniform-like black dagger mark (cf. H. elevatus roraima, Fig. 23). Keith Willmott (pers. comm.) reports: "I found many mislabeled specimens in Schmidt-Mumm's collection, since, at least the specimens examined, were pinned in the bottom half of a hinged box with the labels separate in the top half. It seems specimens frequently got moved around in the bottom half without their labels also being moved. I have to say this looks like a pretty unusual 'elevatus' variant, and some kind of hybrid origin seems perhaps more likely, but since I would say the label data are highly unreliable, we essentially don't know where it was collected at all. So, difficult to speculate further about a hybrid origin." If the locality data are incorrect, it is possible that the specimen is a H. elevatus x H. pardalinus.

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FIGURE 3. Heliconius elevatus pseudocupidineus Neustetter, 1931 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 3. Heliconius elevatus pseudocupidineus Neustetter, 1931 (dorsal; ventral). Peru: Loreto, Balsapuerto, 220m. Feb. 1919, leg. G. G. Klug. (Image source http://www.tolweb.org/, images # 17245 and #17249). H. elevatus is a member of the silvaniform clade (as recognized by Eltringham, 1916), and is sister taxon to H. pardalinus Bates, 1862. Note that H. elevatus bears a yellow subcostal stripe on the HWV, while in rayed H. melpomene forms, the corresponding stripe is orange or red. Also note the tendency for the widened proximal end of the red HW rays to exhibit a v-shape, while in H. melpomene, they are flat "nail heads."

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FIGURE 10. H. numata arcuella Druce, 1874 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 10. H. numata arcuella Druce, 1874 (dorsal, ventral). Peru: Río Ucayali, Pucallpa, 200 m. (image source: https://cliniquevetodax.com/Heliconius/pages/numata%20arcuella.html).

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FIGURE 19 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 19. Hybrid #13 (dorsal, ventral). Specimen (in UFRJ) data poor: no date or collector indicated, locality either Obidos or Benjamin Constant, Brazil (approximately 800 miles apart!—see also hybrid #9). Mallet et al. (2007) interpreted this specimen as a F1 hybrid between "H. hecale fortunatus" Weymer, 1883 and H. melpomene melpomene (Fig. 6). The name fortunatus is a junior synonym of H. pardalinus radiosus (Lamas, 2004). (Fig. 20). If from Benjamin Constant, it would be unlikely to be related to H. melpomene melpomene, which does not occur there. The dentate HW rays suggest that this is a form of H. pardalinus, or possibly a pardalinus x elevatus cross.

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FIGURE 2. Heliconius melpomene aglaope C. Felder & R. Felder, 1862 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 2. Heliconius melpomene aglaope C. Felder & R. Felder, 1862 (dorsal; ventral). Peru: Loreto, Contamana, 170 m. 26 May 2002. leg. C. Peña. (Image source: http://www.tolweb.org/, images #17224 and #17230). This specimen exhibits a "typical" phenotype of a geographical race with the red forewing "dennis" and hindwing "ray" pattern.

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FIGURE 18. H in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 18. H. elevatus bari Oberthür, 1902 (dorsal, ventral). French Guiana: Montagne de Kaw, pk 29.2 (image source: https://cliniquevetodax.com/Heliconius/pages/elevatus%20bari.html)

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FIGURE 9. H. melpomene meriana Turner, 1967 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 9. H. melpomene meriana Turner, 1967 "typical" form (dorsal, ventral). French Guiana. (Image source: https:/ /cliniquevetodax.com/Heliconius/pages/melpomene%20meriana.html).

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FIGURE 20. H. pardalinus radiosus Butler, 1873 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 20. H. pardalinus radiosus Butler, 1873 (dorsal). Brazil: Amazonas, Villa Bella [Bolivia:Pando?]. Neukirchen coll. (FLMNH). (image source: https://cliniquevetodax.com/Heliconius/pages/pardalinus%20radiosus.html)

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FIGURE 17. H. melpomene thelxiopeia Staudinger, 1897 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 17. H. melpomene thelxiopeia Staudinger, 1897 (dorsal, ventral). French Guiana: Rivière Comte, leg. C. Chazal. Note the reduction of ray elements on the HWV, and rounded proximal edges of yellow spots on FW (image source: https://cliniquevetodax.com/Heliconius/pages/melpomene%20thelxiopeia.html).

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FIGURE 14. H. melpomene malleti Lamas, 1988 in Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae)

FIGURE 14. H. melpomene malleti Lamas, 1988 (dorsal, ventral). Peru: Loreto. (image source: https:// cliniquevetodax.com/Heliconius/pages/melpomene%20maletti.html)

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