“…Additional muscle activity means that whole-animal metabolic costs (rate of O 2 consumption, V O2 ) should also increase (Ellerby et al, 2003), assuming that additional costs are met aerobically. Accordingly, increased metabolic costs were found in birds (Bamford and Maloiy, 1980;Ellerby et al, 2003;Rubenson et al, 2006;Snyder and Carello, 2008;Warncke et al, 1988), humans (Dill, 1965;Margaria et al, 1963;Minetti et al, 2002;Yousef et al, 1972), a range of quadrupeds (Armstrong et al, 1983;Bedford et al, 1979;Brooks and White, 1978;Chassin et al, 1976;Clapperton, 1964;Cohen et al, 1978;Eaton et al, 1995;Farley and Emshwiller, 1996;McDonough et al, 2002;Raab et al, 1976;Robbins et al, 1979;Snyder and Carello, 2008;Taylor et al, 1972;White and Yousef, 1978;Wickler et al, 2000;Wickler et al, 2003;Wunder and Morrison, 1974;Yousef et al, 1972) and invertebrates (Full and Tullis, 1990;Tullis and Andrus, 2011) when ascending gradients. Although there are notable exceptions -the marabou stork Leptoptilos crumeniferus (Bamford and Maloiy, 1980), cockroach Gromphadorhina portento (Herreid et al, 1981) and ant Camponotus sp.…”