The webbed feet of waterbirds are morphologically diverse and classified into four types: the palmate foot, semipalmate foot, totipalmate foot, and lobate foot. to understand the developmental mechanisms underlying this morphological diversity, we conducted a series of comparative analyses. Ancestral state reconstruction based on phylogeny assumed that the lobate feet possessed by the common coot and little grebe arose independently, perhaps through distinct developmental mechanisms. Gremlin1, which encodes a bone morphogenetic protein (BMp) antagonist and inhibits interdigital cell death (icD) in the foot plate of avian embryos, remained expressed in the interdigital tissues of webbed feet in the duck, common coot, little grebe, and great cormorant. Differences in Gremlin1 expression pattern and proliferating cell distribution pattern in the toe tissues of the common coot and little grebe support the convergent evolution of lobate feet. in the totipalmate-footed great cormorant, Gremlin1 was expressed in all interdigital tissues at St. 31, but its expression disappeared except along the toes by St. 33. The webbing of the cormorant's totipalmate foot and duck's palmate foot may have risen from distinct developmental mechanisms. Waterbirds usually have webbed feet for foot-based propulsion. Webbed feet can be morphologically classified into four types: palmate, semipalmate, totipalmate, and lobate. Palmate feet are the most common type of webbed feet in waterbirds, where three front-facing toes (toes II, III, and IV) are entirely connected by webbing 1,2 (Fig. 1A). Semipalmate feet have partial webbing at the base of the three front-facing toes (Fig. 1A). Totipalmate feet have a hallux connected by webbing to front-facing toe II, and toes II, III, and IV are entirely connected by webbing (Fig. 1A). The four toes of lobate feet are separated from each other, but each toe has leaf-like membranes (lobes) along the edges that produce propulsion in water 3. Webbed feet are observed in five clades of modern birds: (1) order Anseriformes including palmate-footed ducks and geese and semipalmate-footed magpie geese 1,4,5 ; (2) order Gruiformes including lobate-footed sungrebes and coots 1,4,6 ; (3) the clade composed of order Phoenicopteriformes including the palmate-footed flamingos and order Podicipediformes including the lobate-footed grebes 1,4,7 ; (4) order Charadriiformes including the palmate-footed gulls, semipalmate-footed shorebirds, and lobate-footed genus Phalaropes in the family Scolopacidae 1,4,8 ; (5) the large clade including the orders Eurypygiformes, Phaethontiformes, Gaviiformes, Sphenisciformes, Procellariiformes, Ciconiiformes, Suliformes, and Pelecaniformes with palmate (loons, penguins, and shearwaters) or totipalmate feet (Phaethontiformes and Suliformes) 1,4,9. Webbed feet were acquired independently at least fourteen times in modern birds (Fig. 1B). To fully understand the origins and evolutionary history of webbed feet in birds, it is important to clarify the developmental mechanisms underlying w...