Massive, solidified bone in the wing of a volant courting bird.

One pervasive morphological feature of tetrapods is the pipe-like, often marrow-filled, structure of the limb or long bones. This 'hollow' form maximizes flexural strength and stiffness with the minimum amount of bony material, and is exemplified by truly hollow (air-filled), or pneumatic, humeri in many modern birds. High-resolution microCT scans of the wings of two male club-winged manakins (Machaeropterus deliciosus) uncovered a notable exception to the hollow-tube rule in terrestrial vertebrates; males exhibited solidified ulnae more than three times the volume of birds of comparable body size, with significantly higher tissue mineral densities. The humeri exhibited similar (but less extreme) modifications. Each of the observed osteological modifications increases the overall mass of the bone, running counter to pervasive weight-reducing optimizations for flight in birds. The club-winged manakin is named for a pair of unique wing feathers found in adult males; these enlarged feathers attach directly to the ulna and resonate to produce a distinctive sound used in courtship displays. Given that the observed modifications probably assist in sound production, the club-winged manakin represents a case in which sexual selection by female choice has generated an ecologically 'costly' forelimb morphology, unique in being specialized for sound production at a presumed cost in flight efficiency.