Fig. 1. (A). Relative positions of the anterior and posterior spiracles in dung beetles. (B) Collection localities of the four species of dung beetles used in this study indicating their separated distributions on the subcontinent. x Pachylomerus femoralis; x Scarabaeus gariepinus; x S. striatum; x Circellium bacchus. Climate graphs indicate rainfall in mm (y1 axis, solid bars), and temperature in °C (y2 axis, mean maximum, unbroken line; mean minimum, broken line). Ab, abdominal segment.