Willow ptarmigan complicated changes in plumage several times a year and the differing colour and pattern of the summer plumage: The willow ptarmigan often hybridises with the black grouse ("Tetrao tetrix") and the hazel grouse ("Tetrastes bonasia") and occasionally with the western capercaillie ("Tetrao urogallus"), the spruce grouse ("Falcipennis canadensis") and the rock ptarmigan ("Lagopus muta"). During the Pleistocene, the willow ptarmigan widely occurred in continental Europe. Authors who recognize paleosubspecies have named the Pleistocene willow ptarmigan "L. l. noaillensis" (though the older name "medius" might be the correct one). These marginally different birds are said to have gradually changed from the
Snow camouflage snow camouflage patterns. Charles Darwin mentioned the white winter coloration of the ptarmigan in his 1859 "Origin of Species": The white protective coloration of arctic animals was noted by an early student of camouflage, the naturalist Alfred Russel Wallace, in his 1889 book "Darwinism"; he listed the polar bear, the American polar hare, the snowy owl and the gyr falcon as remaining white all year, while the arctic fox, arctic hare, ermine and ptarmigan change their colour, and observed "the obvious explanation", that it was for concealment, at a time when Darwinism was at a low ebb. Later zoologists such