Identification
The rarely seen nominate form of H. axyridis, featuring a "reversed" pattern of orange-red markings on a black background.
Range
Rare in North America, native to Siberia and northern Asia; occurrence probably limited to genetic backcrossing rather than intentional introduction of this form. Absent from much of the introduced range of the species, including western Europe (J. Bailey pers. comms. and also see Roy et. al. 2015).
Print References
Hosino, Y. 1940. Genetical studies on the pattern types the lady-bird beetle, Harmonia axyridis Pallas. Journal of Genetics 40: 215–228.
Majerus et. al, "Guide to Ladybirds of the British Isles", Field Studies Council 2006.
Michie, L.J., F. Mallard, M.E.N. Majerus & F.M. Jiggins. 2010. Melanic through nature or nurture: genetic polymorphism and phenotypic plasticity in
Harmonia axyridis. Journal of Evolutionary Biology 23: 1699–1707. (
Full Text)
Komai, T. 1956. Genetics of lady beetles. Advances in Genetics 8: 155–188.
Komai, T., M. Chino & Y. Hosino. 1950. Contributions to the evolutionary genetics of the lady-beetle, Harmonia .1. Geographic and temporal variations in the relative frequencies of the elytral pattern types and in the frequency of elytral ridge. Genetics 35: 589–601.
Roy, H.E., & P.M.J. Brown. 2015 Ten years of invasion:
Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in Britain. Ecological Entomology 40(4): 336-348. (
Full Text)
Tan, C.C. 1946. Mosaic dominance in the inheritance of color patterns in the lady-bird beetle, Harmonia axyridis. Genetics 31: 195–210.
Tan, C.C. & J.C. Li. 1934. Inheritance of the elytral color patterns of the lady-bird beetle, Harmonia axyridis Pallas. The American Naturalist 68: 252–265.
Ueno, H. 1994. Intraspecific variation of P2 value in a coccinellid beetle, Harmonia axyridis. Journal of Ethology 12: 169–174.
Wang, S., J.P. Michaud, R.Z. Zhang, F. Zhang & S. Liu. 2009. Seasonal cycles of assortative mating and reproductive behaviour in polymorphic populations of Harmonia axyridis in China. Ecological Entomology 34: 483–491.