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      Morphological evolution in Hyles Hübner, 1819 hawkmoths (Lepidoptera, Sphingidae): reconstructing the ancestral Hyles habitus

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      Nota Lepidopterologica

      Pensoft Publishers

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          Abstract

          Molecular phylogenetic studies suggest that similar wing and body patterns in the hawkmoth genus Hyles Hübner, [1819] do not necessarily reflect a close phylogenetic relationship. To improve our understanding of morphological evolution in these organisms, 75 characters derived from the external adult morphology are explicitly coded and analysed in a maximum parsimony cladistic framework. The results corroborate the hypothesis that wing and body patterns have indeed reappeared in different parts of the phylogeny but the underlying genetic mechanism remains to be determined. By reconstructing the suite of ancestral states of the morphological characters using Bayesian inference, we derived an approximation of the appearance of the proto-Hyles species. The overall habitus of this moth does not display a combination of characters found in any extant Hyles species. Rather, the forewings are most like those of members of the Hyles euphorbiae-complex but with better developed antemedial and postmedial lines, the hindwings are typical Hyles, and the abdominal pattern most closely resembles that of Hyles euphorbiarum (Guérin-Méneville & Percheron, 1835), but with one fewer pairs of black subdorsal patches. Within the context of the subtribe Choerocampina and Sphingidae more generally, the proto-Hyles reconstruction does not resemble any other species apart from Rhodafra opheltes (Cramer, 1780), but this appears to be another instance of convergent pattern expression.

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          Most cited references 7

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          Logical basis for morphological characters in phylogenetics

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            A revised molecular phylogeny of the globally distributed hawkmoth genus Hyles (Lepidoptera: Sphingidae), based on mitochondrial and nuclear DNA sequences.

            The hawkmoth genus Hyles comprises some 29 species with a global distribution. In this study, we augment the previous taxon sampling with more species and add sequences from a nuclear gene to produce a refined phylogenetic hypothesis. A total evidence reconstruction based on Bayesian analysis of the combined mitochondrial (COI, t-RNA-Leu, COII; 2284 bp) and nuclear (EF1alpha; 773 bp) sequences is discussed and compared with the results from separate analyses of the two genes. The total evidence phylogeny corroborates many of the phylogenetic relationships previously postulated within the genus. In addition, the hitherto unsampled enigmatic species Hyles biguttata from Madagascar appears as sister group to Hyles livornicoides from Australia, although support for the relationship is relatively weak. The high level of differentiation of Hyles perkinsi from H. calida (both Hawaii), and the status of these two as sister species, is corroborated by both sources of sequence data. However, their phylogenetic position when mt DNA sequences alone are considered differs markedly from that under total evidence. The previously postulated relationships within the Hyles euphorbiae complex (HEC) s.s. are largely corroborated, but H. dahlii is now more closely related and the HEC s.l. is redefined to include H. zygophylli and H. stroehlei (two species that had not been studied previously using molecular data) and to exclude H. siehei and H. hippophaes. The nuclear sequences alone are insufficiently variable to fully resolve all lineages and the phylogeny suggests that nuclear gene swapping and incomplete lineage sorting have occurred implying recent divergence. The results from the total evidence analysis provide a phylogenetic hypothesis that both corroborates and complements the previous biogeographic scenario, and provides new insights into the origins of several of the included taxa.
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              A molecular phylogeny of the hawkmoth genus Hyles (Lepidoptera: Sphingidae, Macroglossinae).

              The hawkmoth genus Hyles is one of 15 genera in the subtribe Choerocampina of the subfamily Macroglossinae. Due to a remarkable uniformity, morphological characters usually used to identify and classify Lepidoptera at the species level cannot be used in this genus. Instead, we used DNA sequences comprising about 2300 bp derived from the mitochondrial genes COX I, COX II, and tRNA-leucine to elucidate the phylogeny of Hyles. The results corroborate the monophyly of Hyles but conflict with previous internal classifications of the genus based on morphology. Hyles seems to have evolved in the Neotropics during the Oligocene/Eocene epochs and the molecular data (which evolved clock-like) confirm the hypothesis that it is a very young genus that radiated on a global scale rather quickly. We hypothesize its sister group to be one of the genera Deilephila, Theretra or Xylophanes. The Nearctic may have been colonized rapidly by Hyles once the land bridge formed during the Pliocene, since within this same Epoch, the invasion of the Palaearctic appears to have proceeded from the East, via the Bering route. The colonization of Australia appears to have occurred rather early in Hyles radiation, although the route is not clear. We propose that the radiation of the Hyles euphorbiae-complex s. str. (HEC) occurred as recently as the Pliocene/Pleistocene boundary and that its roots can still be reconstructed in Asia. Hyles dahlii is closely related to the HEC, but a sister group relationship to the HEC s. str. cannot be corroborated unequivocally. HEC population ranges appear to have tracked climate oscillations during the Pleistocene Ice Ages, resulting in hybridization around the Mediterranean Sea as they repeatedly intermingled. Comparison of the phylogeny with food plant affiliations leads us to hypothesize that Euphorbia monophagy evolved at least two times independently within Hyles.
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                Author and article information

                Journal
                Nota Lepidopterologica
                NL
                Pensoft Publishers
                2367-5365
                0342-7536
                July 31 2020
                July 31 2020
                : 43
                : 181-210
                Article
                10.3897/nl.43.49512
                © 2020

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