We propose that the ancestors of bats were small, nocturnal, sylvatic
gliders that used echolocation for general orientation. Their echoloca
tion calls were short, low intensify, broadband clicks, which translat
ed into a very short operational range. In the lineage that gave rise
to bats, a switch to stronger, tonal signals permitted the use of echo
location to detect, track, and assess flying insects in subcanopy sett
ings. We propose that these animals hunted from perches and used echol
ocation to detect, track, and assess flying insects, which they attack
ed while gliding. In this way, the perfection of echolocation for hunt
ing preceded the appearance of napping night, which marked the emergen
ce of bats. Flapping night had appeared by the Eocene when at least ei
ght families are known from the fossil record. Stronger signals and ad
aptations to minimize self-deafening were central to the perfection of
echolocation for locating flying prey. Echolocation constituted a key
innovation that permitted the evolution and radiation of bats. At the
same time, however, its short effective range imposed a major constra
int on the size of bats. This constraint is associated with flight spe
ed and the very small time intervals from detection of, and contact wi
th a flying target. Gleaning and high duty cycle echolocation are two
derived approaches to hunting prey in cluttered situations, places whe
re echoes from background and other objects arrive before or at the sa
me time as echoes from prey. Both had appeared by the Eocene.