Interspecific mutualisms are regarded as having evolved from antagonis
tic or commensalistic interactions, with most mutualisms remaining fac
ultative but some having coevolved into obligate reciprocal dependency
(1-4). Underlying mutualism is an intrinsic conflict between the parti
es, in that each is under selection for increased exploitation of the
other(3-7). Theoretical models suggest that this conflict is a source
of evolutionary instability, and that evolution of 'cheating' by one p
arty may lead to reciprocal extinction(4,6,7). Here we present phyloge
netic evidence for reversal of an obligate mutualism: within the yucca
moth complex, distinct cheater species derived from obligate pollinat
ors inflict a heavy cost on their yucca hosts by laying their eggs but
not pollinating the yucca. Phylogenetic data show the cheaters to hav
e existed for a long time. Coexisting pollinators and cheaters are not
sister taxa, supporting predictions that evolution of cheating within
a single pollinator is evolutionarily unstable. Several lines of evid
ence support a hypothesis that host shifts preceded the reversal of ob
ligate mutualism. Host or partner shifts is a mechanism that can provi
de a route of evolutionary escape among obligate mutualists in general
.