“…The basic, non-flight related, functions of bird body feathers are twofold: First, by varying the number of feathers per surface area, the amount of trapped air, and hence water resistance and thermo-conductivity, can be controlled (e.g., Middleton 1986;Swanson 1991;Fernando Novoa, Bozinovic & Rosenmann 1994;Cooper 2002;Williams, Hagelin & Kooyman 2015). Second, by varying the composition, microoptical and microstructural properties of plumage (i.e., color, relative densities of different feather types, morphologies of body feathers), water repellency, and insulatory functions can also be controlled (e.g., Rijke 1970;Middleton 1986;Wolf & Walsberg 2000;Ward et al 2007;Broggi et al 2011;Rijke & Jesser 2011;Pap et al 2015;Williams, Hagelin & Kooyman 2015;Koskenpato et al 2016). These different functions are immediately evident in different species; the Emperor Penguin (Aptenodytes forsteri), for example, has a particular spatial pattern and ratio of different feather types, including body and attached after-feathers, filoplumes, and downy feathers that make its body covering uniquely insulative and suited to its lifestyle (Williams, Hagelin & Kooyman 2015), while diving Great Cormorants (Phalacrocorax carbo) balance body feather structures between waterproofing and wettability to fulfil requirements of insulation and reduced buoyancy during diving (Rijke 1968;Gr emillet et al 2005).…”