A mechanistic approach to understanding the colour polymorphism in black sparrowhawks (Accipiter melanoleucus)

Abstract

Species that exhibit large variation in phenotypic traits, are commonly considered to have a stronger evolutionary potential. However, how they are capable to maintain polymorphisms remains a fundamental problem in evolutionary biology. Colour polymorphic species provide ideal study systems to explore the processes that lead to variation maintenance. About 3.5% of all bird species are colour polymorphic, but it is an especially common phenomenon in Accipitridae (22%), which indicates that it has an adaptive function in this bird group and makes them ideal model systems to study evolutionary processes. The black sparrowhawk (Accipiter melanoleucus) occurs in two discrete colour variants: dark and light. The two morphs differ in the expression of white and black feathers on the breast, belly and underwing coverts. The morph has been associated with ambient light-dependent foraging success and activity behaviour: Dark morphs forage more and have higher foraging success under low light conditions whereas light morphs forage independently of light levels but are better foragers under bright light conditions. This is hypothesized to be due to a crypsis advantage for the morphs under these conditions. During the winter breeding period, the predominating low light conditions on the Cape Peninsula (Western Cape, South Africa) could create an advantage for dark morphs, which should result in their higher survival and higher breeding success. However, this is not the case and only when the two morphs come together to breed, there is a fitness difference: mixed-morph pairs (that consist of a dark and a light morph) have higher breeding success than like-morph pairs (that consist of the same morph) and offspring of mixed-morphs have higher survival rates. This higher success of mixed-morph pairs is hypothesized to be due to emergent pair-level properties with the two morphs being able to expand the hunting niche as a pair. This ‘complementarity hypothesis' is based on previous research conducted on the study system. The aim of this PhD was to explore the mechanistic background of colour polymorphism maintenance in the black sparrowhawk. (i) I performed an experiment in which I test whether there is a morph- and ambient light-dependent crypsis advantage in the black sparrowhawk. I measure the reaction time of feral pigeons towards a simulated hawk attack but did not find indication of such an effect as pigeons reacted the same towards the two morphs. In line with the complementarity hypothesis, (ii) I found that mixed-morph parents provide food more consistently to the nest than like-morph parents. This results in a more predictable food supply for nestlings and buffers against long periods of malnourishment. However, (iii) I was unable to determine the mechanistic link between food supply and higher survival: nestlings of mixedand like-morph pairs had the same levels of innate immune function. Thus, an improved innate immune function in nestlings of mixed-morph pairs is unlikely associated with their higher survival rates. (iv) I performed individual-based model simulations which incorporate multiple key fitness parameters and found that complementarity – in combination with morphdependent seasonality-associated fitness effects – explains the stable colour morph equilibrium in this population. I conclude that emergent pair-level properties which arise due to the complementary nature of the two morphs play an important role in maintaining polymorphism in this species. Complementarity might not only be restricted to colour polymorphic species but could be present in other polymorphic traits that allow parents to behaviourally complement each other when raising their young

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