Phylogeny and chemicals ecology of Hyles hawkmoths (Lepidoptera, Sphingidae) with special reference to the Hyles euphorbiae complex
Final Report Abstract
Total evidence analyses of nuclear and mitochondrial sequences of the genus Hyles, including several additional species, elucidated main aspects of Hyles phylogeny, including structure within the Hyles euphorbiae complex (HEC) s.l.. Biogeographic ancestral range analyses corroborate the genus Hyles to be of Neotropical origin, radiating to Southern Hemisphere Madagascar and Australia from the Neotropics. The Northern Hemisphere was then colonised from the Neotropics, the routes to Hawaii and the Palaearctic being via the Nearctic, thus requiring only a single evolutionary transition from the tropical zone to the temperate zone. The Nearctic would have been independently colonised three times during Hyles evolution: twice from the Neotropics (H. lineata and the ancestor of the temperate clade) and a third time by a H. gallii clade. Our data indicate that this last colonisation event is very recent, at the earliest in the Late Pleistocene. We propose an Eastern and/or Central Palaearctic origin of the Hyles radiation in the Palaearctic, once this region had been colonized from Nearctic stock. The analyses infer two colonisation events of the genus that must have been the result of extremely rapid intercontinental range expansions/dispersals in rather recent Hyles evolution: Palaearctic H. livornica into Southern Africa and Palaearctic H. gallii into the Nearctic, underlining that Hyles hawkmoths are strong dispersers. Microsatellite data of over 800 individuals indicate partition of the entire HEC into two clusters roughly corresponding to the two morphologically postulated main lineages Hyles euphorbiae in Asia and Europe and H. tithymali in Macaronesia. A very small FST value demonstrates the low nuclear genetic divergence observed between the two clusters. The HEC exhibits a strong signal of neighbour-mating across its entire range, i.e. isolation-by-distance appears to be an important factor shaping the population structure. Augmented sequencing of mitochondrial haplotypes also corroborated the separation of the HEC into two main partitions: H. tithymali lineages in Macaronesia, Maghreb, Southern Italy, the Aegean Islands and Yemen and H. euphorbiae lineages on the Iberian and Balkan Peninsula, Central and Eastern Europe and the Near East. However, main parts of Europe and the Near East (the range of H. euphorbiae) are sympatrically occupied by three distinct lineages. The euphorbiae lineage is most abundant and distributed throughout this range with further extra-limital occurrences in Southern Italy, on Malta, Pantelleria and the Cape Verde Islands. The 'enigmatica' lineage is most abundant in Eastern and Central Europe and the North Eastern parts of the Apennine peninsula but rare in Western Europe. The very distinct 'melitensis' lineage was now also detected on Sicily and in on mainland Europe, although it had first been found to be endemic to Malta. Yemen, Northern Africa (Maghreb) and Macaronesia (the range of H. tithymali) are predominantly occupied by a single lineage, tithymali, which has also been detected near Rome and on Pantelleria. The cretica and 'italica' clusters are closely related to tithymali. The former is confined to Crete and the Dodecanese Islands (the range of H. cretica). The latter lineage, 'italica', has its main distribution centre in the Apennine Peninsula and on Sicily and Pantelleria. Comparison of genetic markers demonstrates the incongruence between mitochondrial and nuclear sources of data in many areas, indicating a high amount of genomic introgression. Larval morphotype distribution also suggests high levels of hybridisation. The hypothesis that the HEC exhibits a very wide hybridisation zone is thus corroborated by the total evidence of morphology, mitochondrial and nuclear genetic markers. The area covered by the hybridisation zone is extended to include southern Europe, Northern Africa and Yemen. By significantly raising the sample size and including new localities the course of the project led to the discovery of an unexpected, new derived mitochondrial lineage on Pantelleria and in Tunisia which appears to be endemic to the Central Mediterranean, similar to what has been hypothesized for 'melitensis' on Malta. In addition, with the significant rise of samples from Central and Eastern Europe, we could also show that this 'melitensis' lineage is widely distributed among the lineages euphorbiae and 'enigmatica' on the mainland and thus not endemic to Malta and possibly not of Maltese origin but rather could have dispersed to the island in a similar way as hypothesized for the euphorbiae lineage.
Publications
- (2009) A revised molecular phylogeny of the globally distributed hawkmoth genus Hyles (Lepidoptera: Sphingidae), based on mitochondrial and nuclear DNA sequences. Molecular Phylogenetics and Evolution 52, 852-865
Hundsdoerfer, A.K., Rubinoff, D., Attié, M., Wink, M. & Kitching, I.J.
- (2010) Eleven hawkmoth microsatellite loci of Canary Island Hyles tithymali (Lepidoptera). Conservation Genetics Resources 2, 241–244
Hundsdoerfer, A.K., Sanetra, M., Corbeil, D. & Stuckas, H.
- (2011) Larval pattern morphotypes in the Western Palaearctic Hyles euphorbiae complex (Lepidoptera: Sphingidae, Macroglossinae). Insect Systematics & Evolution 42, 41-86
Hundsdoerfer, A.K., Mende, M.B., Harbich, H., Pittaway, A.R., & Kitching, I.J.
- (2011) Taxonomy, phylogeography and climate relations of the Western Palaearctic spurge hawkmoth (Lepidoptera, Sphingidae, Macroglossinae). Zoologica Scripta 40(4), 403-417
Hundsdoerfer, A.K., Mende, M.B., Kitching, I.J. & Cordellier, M.
