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 3K8X2 Linnaeus 1758 [371,470,151,177] Insecta Nymphalidae Heliconius GBIF Animalia Lepidoptera 68 69 Arthropoda species erato  For most of the 20th century,  H. himerawas viewed as a race of  H. erato( Eltringham 1916; Brown 1979), and from a phylogenetic perspective,  H. himerais nested among various races of a paraphyletic  H. erato( Brower 1994b; Hines et al.2011; Supple et al.2015; Van Belleghem et al.2017). These two taxa are parapatric in southern Ecuador, where  H. erato cyrbiaprefers wetter, lower forest, and  H. himeraoccurs in higher, drier  Acaciascrub, however they abut in a narrow hybrid zone ( Jiggins et al. 1996). Descimon & Mast de Maeght (1984) suggested a "semispecies" relationship between the two, due to a deficit of observed hybrid specimens from numbers expected if the parental forms were mating at random. Based on this observation, Mallet (1993, p.245)opined, "it is more sensible to regard  H. himeraas a good species." Mallet (1995)proposed the "genealogical cluster concept" of species to accommodate taxa such as  H. himera, which while still hybridizing at appreciable frequency with its sister taxon appears to have reached a tipping point at which homogenizing gene flow does not occur. We will return to Mallet's species concept in the discussion. Mallet et al. (2007) listed a series of 52 hybrid specimens between  H. erato cyrbiaand  H. himera(three exemplars of which are shown below: Figs. 158–160). The genetics of wing pattern inheritance were experimentally worked out by Jiggins et al.(1996), and there is no doubt that these specimens are legitimate hybrids. Thus, hybrids #101-152 are given the following collective scores: Identity: 1.0 Authenticity: 1.0 Overall reliability: 1.0   FIGURE 158.Hybrid #111 (dorsal, ventral).  H. erato cyrbia(Fig. 156) x  H. himera(Fig. 157) F1 (dorsal, ventral). Ecuador: El Oro, Guayquichuma site 4, 1993 (Neukirchen collection, FLMNH). The yellow forewing bands are replaced with red, but otherwise the specimen looks like  H. himera.   FIGURE 159.Hybrid #110 (dorsal, ventral).  H erato cyrbia(Fig. 156) x  H. himera(Fig. 157) backcross to  H. himera.Ecuador: El Oro, Guayquichuma site 4, 1993. leg. S. Attal (Neukirchen collection, FLMNH). There is a faint pinkish edge on the distal margin of the yellow forewing band.   FIGURE 160.Hybrid #117(dorsal, ventral).  H erato cyrbia(Fig. 156) x  H. himera(Fig. 157) backcross to  H. erato.Ecuador: El Oro, Guayquichuma site 4, 1993. leg. S. Attal (Neukirchen collection, FLMNH). Both H. himera'sred HW band and the yellow HWV stripe of  H. erato cyrbiaare expressed in this individual.   FIGURE 161.  Heliconius erato favorinusHopffer, 1874(dorsal, ventral). Peru: Huanuco, Tingo Maria. (image source: https://cliniquevetodax.com/ Heliconius/pages/ erato%20 favorinus.html)   FIGURE 162.Hybrid #153 (dorsal). Peru: San Martín, Rodriguez de Mendoza, 1500m. Nov. 1984, leg. F. König, König collection (NMW). Austrian Fritz König (d. 2102) lived and collected in Peru from 1954-1980's(?), and his collection was donated to Naturhistorisches Museum, Wien after his death. König (1986) reported this specimen as an intraspecific hybrid (viewing  H. himeraas a race of  H. erato). Mallet et al.(2007) interpret it as an  H. erato favorinus(Fig. 161) x  H. himera(Fig. 157) F1. Given the locality and what is known about inheritance of the various wing pattern elements in these hybrids (Jiggins et al.1996), this interpretation seems plausible. Identity: 1.0 Authenticity: 1.0 Overall reliability: 1.0   FIGURE 163.Hybrid #154 (dorsal, ventral). Peru: San Martín, Rodriguez de Mendoza, 1994, leg. M. Bueche. Neukirchen coll. (FLMNH). Interpreted by Mallet et al.(2007) as an  H. erato favorinus(Fig. 161) x  H. himera(Fig. 157) backcross to  H. erato—presumably based on the locality data. The distal margin of the FW red band is not like either of the putative parental forms, and the truncation of the yellow HW band is also not evident in other putative  H. eratox  H. himerahybrids. Further, this specimen is very similar to the holotype of  Heliconius amatusStaudinger, 1897(see http:// www.butterfliesofamerica.com/L/ih/ heliconius0490_i.htm), which is considered to be a  H. erato phyllis(Fabricius, 1775)x  H. erato venustusSalvin, 1871backcross to  H. erato phyllis. As noted for the specimen illustrated in Fig. 42, Martin Bueche runs a butterfly house in France. It is very probable that that the locality data are false, and it may well have been reared in captivity. Identity: 0.5 Authenticity: 0.2 Overall reliability: 0.1 Hybrid #155 (not illustrated in Mallet et al.2007). Peru: San Martín, Rodriguez de Mendoza-Omia km 11, 1986, leg. J. Mallet (Mallet collection). Interpreted by Mallet et al.(2007) as  H. himera(fig. 157) x  H. erato favorinus(fig. 161). This specimen cannot be independently evaluated from the evidence presented.  Habeas corpus! Identity: 0. Authenticity: 1.0 Overall reliability: 0.0   FIGURE 164.Hybrid #156 (dorsal). Peru: San Martín, Rodriguez de Mendoza-Omia km 11, ~1986. König collection (NMW). Interpreted by Mallet et al.(2007) as  H. himera(Fig. 157) x  H. erato favorinus(Fig. 161) backcross to  H. erato. The locality data of this specimen are the same as Hybrid #155. König (1986) does not mention this specimen, but it is not as dramatic a hybrid as Hybrid #153. The shape of the FW red band is more consistent with those of the putative parental forms than is that of Hybrid #154. Identity: 0.9 Authenticity: 0.8 Overall reliability: 0.72   FIGURE 165.Hybrid #15 (dorsal). 7. Peru: Amazonas: Aramango, <1990. König coll. (NMW). Interpreted by Mallet et al.(2007) as a  H. erato lativitta(Fig. 166) x  H. himera(fig. 157) F1. The dennis and ray pattern of this specimen, combined with the position and shape of the yellow FW band and the red transverse basal band on the HW represent a tidy combination of features of the putative parental forms. Mallet (1993) reported finding no hybrids in the putative zone of contact between  H. himeraand  H. erato lativitta(in his collections, the two populations were geographically separated by approximately 10 km). Identity: 1.0 Authenticity: 0.9 Overall reliability: 0.9