Alternative facts: a reconsideration of putatively natural interspecific hybrid specimens in the genus Heliconius (Lepidoptera: Nymphalidae) Brower, Andrew V. Z. Zootaxa 2018 2018-10-12 4499 1 1 87 3K8WQ cydno Doubleday 1847 [379,598,151,178] Insecta Nymphalidae Heliconius GBIF Animalia Lepidoptera 44 56 Arthropoda subSpecies cydno cydno  Identity: 0.8 Authenticity: 0.8 Overall reliability: 0.64   FIGURE 99.  H. cydno lisethaeNeukirchen, 1995(dorsal, ventral). Colombia: Huila, San Agustín. (image source: https:/ /cliniquevetodax.com/ Heliconius/pages/ cydno%20 lisethae)   FIGURE 100.Hybrid #65 (dorsal, ventral). "Nouve Granade" (no additional locality data). Holotype of  H. aventinaOberthür, 1925(NHMUK). Viewed by Ackery & Smiles (1976) as a  H. cydno hermogenesx  H. melpomene melpomenehybrid; viewed by Mallet et al.(2007) as a  H. melpomene"{ euryas}" (Fig. 73) x  H. cydno hermogenes(Fig. 101) F1 (see nomenclatural notes under Hybrid #41).  H. cydno hermogenesoccurs in the upper Magdalena Valley, between the ranges of  H. cydno lisethae(Fig. 99) to the south, and  H. cydno cydno(Fig. 72) to the north.  H. cydno hermogenesis the only  H. cydno race witha double row of marginal/submarginal spots on the FW. This specimen has characters most consistent with that race, other than the brick-red patch distal to the discal cell on the forewing and the absent hourglass mark in the FW discal cell. Note also that the FW spots are elongated into streaks. Given the lack of locality data, this specimen could also possibly represent a hybrid with a red-banded form of  H. heurippa. Identity: 0.4 Authenticity: 0.4 (due to lack of locality data) Overall reliability: 0.16   FIGURE 101.  H. cydno hermogenesHewitson[1858] (dorsal, ventral). Colombia: [Tolima], Rio Chili. (image source: https://cliniquevetodax.com/ Heliconius/pages/ cydno%20 hermogenes). Note that the infrasubspecific named form " lutescens" Kaye, 1916 has yellow, instead of white spots on the forewing.  Putative H. melpomenex  H. cydnohybrids from Chocó, Colombia Hybrid #66 (not illustrated—see plate 35, #4b in Holzinger & Holzinger, 1994. —That figure, quite similar to Hybrid #43, shows a rusty-red distal forewing band on an otherwise rather typical—looking  H. cydnophenotype). "W. Colombia" leg. E. Krüger. Holotype of  H. cydno chioneusf. " eugenius" Niepelt, 1928. NMW(?). Interpreted (presumably on the basis of the locality data) by Mallet et al.(2007) as  H. cydno chioneus? ( Fig. 60) x  H. melpomene vulcanus( Fig. 102) F1 backcross to  H. cydno. The available information on this specimen renders it impossible to interpret. Identity: 0.0 Authenticity: 0.2 Overall reliability: 0.0   FIGURE 102.  H. melpomene vulcanusButler, 1865(dorsal, ventral). Colombia: [Valle del Cauca], Río Dagua. This is one of the most striking forms of  H. melpomene, with a bluish iridescence on the dorsal surface and a fine fringe of white scales on the wing margins. Also note that the transverse yellow HW band is only expressed on the ventral surface. (image source: https://cliniquevetodax.com/ Heliconius/pages/ melpomene%20 vulcanus.html).   FIGURE 103.Hybrid #67 (dorsal). Colombia (W. Colombia), leg. E. Krüger (MZPW). Interpreted (presumably on the basis of the locality data) by Mallet et al.as  H. cydno chioneus(Fig. 60) x  H. melpomene vulcanus(Fig. 102) F1. The HW submarginal band does not reach the margin, which does not reflect the typical pattern in  H. cydno chioneus. The FW band in  H. melpomene vulcanusis compact, well-defined and carmine-red, not the brick-red of the red patch here. The vague locality data on this specimen (label in image is illegible) render it difficult to interpret. It may be a hybrid between some alternative races, or possibly an atavistic mutant. Identity: 0.2 Authenticity: 0.5 Overall reliability: 0.1   FIGURE 104.Hybrid #68 (dorsal, ventral). Colombia: Chocó, Río San Juan, Palestrina. 1988. leg. C. Farrell. Neukirchen Collection (FLMNH). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 105) x  H. melpomene vulcanus(Fig,. 102) F1 backcross to  H. cydno, based on the large white forewing band and completely expressed brown forceps on the HWV. The apparent source of this specimen, as well #69-#71 and #85 and #86, was Christopher Farrell, a commercial butterfly dealer who resided in Bogotá, Colombia in the 1980's (Johnson & Matusik, 1986; see also http:// insectnet.proboards.com/thread/441) (not Clive Farrell, owner of large butterfly houses in the U.S. and the U.K. with many captive-reared  Heliconius). Given the commercial source, and that other specimens in the Neukirchen Collection are also of dubious authenticity, the parsimonious explanation is that this specimen was reared for the trade. Identity: 0.9 Authenticity: 0.1 Overall reliability: 0.09   FIGURE 105.  H. cydno zelindeButler, 1869(dorsal, ventral). Colombia: Cali, Río Dagua [the race occurson the Pacific slope in Valle del Cauca and Chocó, not in the Cauca Valley where Cali is situated]. (image source: https:// cliniquevetodax.com/ Heliconius/pages/ cydno%20 zelinde.html).   FIGURE 106.Hybrid #69 (dorsal, ventral). Colombia: Chocó, Río San Juan, Palestrina. 1988. leg. C. Farrell. Neukirchen Collection (FLMNH). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 105) x  H. melpomene vulcanus(Fig. 102) F1, based on the larger red forewing band and yellowish tint to the anterior band of the brown forceps on the HWV ( H. melpomene vulcanushas a yellow stripe on the HWV). As with Hybrid #68, this specimen likely was reared for the commercial trade. Identity: 0.9 Authenticity: 0.1 Overall reliability: 0.09   FIGURE 107.Hybrid #70 (dorsal, ventral). Colombia: Chocó, Río San Juan, Palestrina. 1988. leg. C. Farrell. Neukirchen Collection (FLMNH). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 105) x  H. melpomene vulcanus(Fig. 102) F1. Specimen is quite similar to Hybrid #69, and interpreted to have the same provenance. Identity: 0.9 Authenticity: 0.1 Overall reliability: 0.09   FIGURE 108.Hybrid #71 (dorsal, ventral). Colombia: Chocó, Río San Juan, Palestrina. 1988. leg. C. Farrell. Neukirchen Collection (FLMNH). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 105) x  H. melpomene vulcanus(Fig. 102) F1, based on the larger red forewing band and partial suppression of the brown forceps on the HWV. Also likely reared for commerce. Identity: 0.9 Authenticity: 0.1 Overall reliability: 0.09   FIGURE 109.Hybrid #85 (dorsal, ventral). Colombia: Valle, Río Calima, near Río Bravo, 1200m. 1988. leg. C. Farrell. Neukirchen Collection (FLMNH). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 105) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno. Other than the slightly wider white FW band and the different locality data, this specimen appears to fit rather well into this series of material "collected" by dealer Farrell. Identity: 0.9 Authenticity: 0.1 Overall reliability: 0.09   FIGURE 110.Hybrid #86 (dorsal, ventral). Colombia: Valle, Río Calima, near Río Bravo, 1200m. 1988. leg. C. Farrell. Neukirchen Collection (FLMNH). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 105) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno. Very similar to Hybrid #69, but with different locality data. This specimen also appears to fit rather well into the series of material "collected" by dealer Farrell. Identity: 0.9 Authenticity: 0.1 Overall reliability: 0.09   FIGURE 111.Hybrid #72 (dorsal, ventral). Colombia: Valle, Cali, 1973. Leg. L. Denhez? Mast de Maeght coll. (RBINS—James Mast de Maeght donated his large butterfly collection to the Royal Belgian Institute of Natural Sciences in 2015). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 105) x  H. melpomene vulcanus(Fig. 102) F1, based on the larger red forewing band and partial suppression of the brown forceps on the HWV. León Denhez (and his son Leoncito?) were commercial collectors based in Cali, whose atypical  Heliconiusmaterial appears not only in Mast de Maeght's collection, but also in the Holzinger and Neukirchen collections. It is likely that the locality data on the label are the source of the material from the dealer, rather than the place where the specimen was collected. This is supported by the high degree of similarity between this specimen and Hybrid #71, leading to Mallet et al.'s interpretation that the two represent crosses of the same races—the endemic race of  H. cydnoin the vicinity of Cali is  H. cydno cydnides(Fig. 112), and there is no sympatric race of  H. melpomenethat occurs there. Once again, in light of the commercial origin of the specimen and the abundance of "hybrids" from this source in the collections of European amateurs, the provenance of the specimen is dubious. Identity: 0.6 (poor locality data) Authenticity: 0.1 Overall reliability: 0.06   FIGURE 112.  H. cydno cydnidesStaudinger, 1885(dorsal, ventral). Colombia: "Valle del Cauca, Río Dagua" [probably incorrect locality data, as this race occursin the Cauca Valley, not on the Pacific slope]. (image source: https:// cliniquevetodax.com/ Heliconius/pages/ cydno%20 cydnides)   FIGURE 113.Hybrid #73 (dorsal, ventral). Colombia: Valle, Buenaventura, Isla Punta Soldado, 30 March(?) 2000. Efraín Henao collection (Universidad Nacional de Colombia, Bogotá). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 112) x  H. melpomene vulcanus(Fig. 102) F1. The specimen is quite similar to Hybrid #72, except with a more ragged outer margin to the FW red band, and a lack of submarginal white spots on the HWV. This specimen was collected by a butterfly biologist, and looks like it actually spent some time flying around before that event. Identity: 0.9 Authenticity: 1.0 Overall reliability: 0.9   FIGURE 114.Hybrid #74 (dorsal, ventral). Colombia: Valle, Río Calima, Estación Agrícola del Bajo Calima, 1979, leg. A. Jaramillo (ICNB). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 112) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. melpomene. The specimen appears to have a shadow yellow stripe on the dorsal HW (apparently absent on the VHW), which is not a characteristic of  H. melpomene vulcanus. It does not look like any of the other Chocó hybrids, and could be an aberrant  H. melpomene. Identity: 0.6 Authenticity: 1.0 Overall reliability: 0.6   FIGURE 115.Hybrid #75 (dorsal, ventral). Colombia: Valle, Cali, 1969. Leg. L. Denhez? Mast de Maeght coll. (RBINS). See provenance notes under Hybrid #72. Interpreted by Mallet et al.(2007) as  H. cydno weymerif. "  weymeri" (Fig. 116) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno, based on the narrow yellow bar on the hindwing. The "hybrid" features are very subtle, and given the likely commercial origin and poor locality data of this specimen, it cannot be regarded as evidentially reliable. Identity: 0.6 (poor locality data) Authenticity: 0.1 Overall reliability: 0.06   FIGURE 116.  H. cydno weymeriStaudinger 1897("typical" form, dorsal, ventral). Colombia: Río Cauca valley. This race is polymorphic (see Fig. 118).   FIGURE 117.  H. cydno weymerif. " gustavi" Staudinger, 1897 (dorsal, ventral). Colombia [no further locality data]. The name gustaviis an intrasubspecific quadrinominal and not available under the ICZN Code (ICZN, 1999). Some " gustavi" individuals have pure black forewings. This specimen is atypical, with white spots on the FW and a partial brown forceps on the VHW. It may be an intraspecific hybrid backcross to another  H. cydnoform (image source: https:// cliniquevetodax.com/ Heliconius/pages/ cydno%20 weymeri).   FIGURE 118.Hybrid #76 (dorsal, ventral). Colombia: Valle, Río Dagua, 1987. Leg. L. Denhez/Gerstner. Neukirchen collection (FLMNH). Interpreted by Mallet et al.(2007) to be a  H. cydno weymerif. " gustavi" (Fig. 117) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno, due to the faint red FW bands. A commercially-obtained specimen from Denhez (see Hybrid #73) via Gerstner (see Hybrid #23). There is no reason to believe that this specimen was collected in the wild. Identity: 0.9 Authenticity: 0.1 Overall reliability: 0.09   FIGURE 119.Hybrid #77 (dorsal, ventral). Colombia: Valle, Río Dagua, (no date). Leg. L. Denhez/Gerstner. Neukirchen collection (FLMNH). Interpreted by Mallet et al.(2007) to be a  H. cydno weymerif. " gustavi" (Fig. 117) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno, due to the faint red FW bands. As for Hybrid #76, there is no reason to believe that this specimen was collected in the wild. Identity: 0.9 Authenticity: 0.1 Overall reliability: 0.09   FIGURE 120.Hybrid #78 (dorsal, ventral). Colombia: Valle, Rio Dagua, 1969. Leg. L. Denhez. NMW (Holzinger collection?). Interpreted by Mallet et al.(2007) as a  H. cydno weymerif. "  weymeri" (Fig. 116) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno, based on the presence of the yellow bar on the HWV only. It is not immediately clear why this specimen could not be an intraspecific hybrid of  H. cydno weymeri(Fig. 116) x  H. cydno cydnides(Fig. 112), which meet in a well-studied hybrid zone in Valle del Cauca (cf. Linares 1997). Holzinger & Holzinger (1968) described several infrasubspecific forms of  H. cydnothat they obtained from Denhez via Gerstner (including forms " denhezi" and "  gerstneri"), now deemed to be hybrids between these two  H. cydnoraces (cf. Lamas & Jiggins 2017). Regardless of its identity, the commercial provenance diminishes the specimen's plausibility as a wild-caught hybrid. Identity: 0.5 Authenticity: 0.1 Overall reliability: 0.05   FIGURE 121.Hybrid #79 (dorsal, ventral). Colombia: Valle, Rio Dagua, 1969. Leg. L. Denhez? Mast de Maeght collection (RBINS). Interpreted by Mallet et al.(2007) as a  H. cydno weymerif. "  weymeri" (Fig. 116) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno, based on the partial expression of the yellow bar on the dorsal and ventral HW. See Hybrid #78 for explanation. Identity: 0.5 Authenticity: 0.1 Overall reliability: 0.05   FIGURE 122.Hybrid #80 (dorsal, ventral). Colombia: Valle, Rio Dagua, 1967. Leg. L. Denhez? Mast de Maeght collection (RBINS). Interpreted by Mallet et al.(2007) as a  H. cydno weymerif. "  weymeri" (Fig. 116) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno, based on the relatively narrow yellow bar on the HWV. See Hybrid #78 for explanation. Identity: 0.5 Authenticity: 0.1 Overall reliability: 0.05   FIGURE 123.Hybrid #81 (dorsal, ventral). Colombia: Valle, Rio Dagua, ~1970. Leg. Gerstner. Mast de Maeght collection (RBINS). Interpreted by Mallet et al.(2007) as a  H. cydno zelinde(Fig. 105) x  H. melpomene vulcanus(Fig. 102) F1 or perhaps backcross to  H. melpomene, based on the broad brown forceps on HWV. See Hybrids #69 and #73. As stated above, Hermann Gerstner was a commercial dealer, and middle man for sellers of novelties. Given its provenance and perfect condition, this specimen is not plausible as a wild-caught hybrid. Identity: 0.8 Authenticity: 0.1 Overall reliability: 0.08   FIGURE 124.Hybrid #82 (dorsal, ventral). Colombia: Valle, Rio Dagua, 1968. Leg. L. Denhez? Mast de Maeght collection (RBINS). Interpreted by Mallet et al.(2007) as a  H. cydno weymerif. "  weymeri" (Fig. 116) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno, based on the partial expression of the yellow bar on the dorsal and ventral HW. See Hybrid #79. Commercial origin as described above. Identity: 0.5 Authenticity: 0.1 Overall reliability: 0.05   FIGURE 125.Hybrid #83 (dorsal, ventral). Colombia: Valle, Rio Dagua, 1966. Leg. L. Denhez? Mast de Maeght collection (RBINS). Interpreted by Mallet et al.(2007) as a  H. cydno weymerif. "  weymeri" (Fig. 116) x  H. melpomene vulcanus(Fig. 102) F1 backcross to  H. cydno, based on the partial expression of the yellow bar on the dorsal and ventral HW. See Hybrid #79. Commercial origin as described above. Identity: 0.5 Authenticity: 0.1 Overall reliability: 0.05   FIGURE 126.Hybrid #84 (dorsal, ventral). Colombia: Valle, Rio Dagua, 1966. Leg. L. Denhez. Mast de Maeght collection (RBINS). Paratype of  H. cydno gerstnerif. pseudoweymeriH. & R. Holzinger, 1968 (an unavailable infrasubspecific quadrinominal). Interpreted by Mallet et al. (2007) as a  H. cydno weymerif. “  weymeri” (Fig. 116) x  H. melpomene vulcanus(Fig. 102) backcross to  H. cydno, as but see #78. Furthermore, likely commercial origin as described above. Indentity: 0.5 Authenticity: 0.1 Overall reliability: 0.05