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Physical gestures are prominent features of many species’ multi- modal displays, yet how evolution incorporates body and leg movements into animal signaling repertoires is unclear. Andro- genic hormones modulate the production of reproductive signals and sexual motor skills in many vertebrates; therefore, one possi- bility is that selection for physical signals drives the evolution of androgenic sensitivity in select neuromotor pathways. We exam- ined this issue in the Bornean rock frog (Staurois parvus, family: Ranidae). Males court females and compete with rivals by per- forming both vocalizations and hind limb gestural signals, called “foot flags.” Foot flagging is a derived display that emerged in the ranids after vocal signaling. Here, we show that administration of testosterone (T) increases foot flagging behavior under seminatural conditions. Moreover, using quantitative PCR, we also find that adult male S. parvus maintain a unique androgenic phenotype, in which androgen receptor (AR) in the hind limb musculature is expressed at levels ∼10× greater than in two other anuran species, which do not produce foot flags (Rana pipiens and Xenopus laevis). Finally, because males of all three of these species solicit mates with calls, we accordingly detect no differences in AR expression in the vocal apparatus (larynx) among taxa. The results show that foot flagging is an androgen-dependent gestural signal, and its emergence is associated with increased androgenic sensitivity within the hind limb musculature. Selection for this novel gestural signal may therefore drive the evolution of increased AR expres- sion in key muscles that control signal production to support adap- tive motor performance.


androgen receptor, testosterone, courtship behavior, signal evolution, frogs





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