Eocene archaeocete whales gave rise to all modern toothed and baleen whales (Odontoceti and Mysticeti) during or near the Eocene-Oligocene transition. Odontocetes have asymmetrical skulls, with asymmetry linked to high-frequency sound production and echolocation. Mysticetes are generally assumed to have symmetrical skulls and lack high-frequency hearing. Here we show that protocetid and basilosaurid archaeocete skulls are distinctly and directionally asymmetrical. Archaeocete asymmetry involves curvature and axial torsion of the cranium, but no telescoping. Cranial asymmetry evolved in Eocene archaeocetes as part of a complex of traits linked to directional hearing (such as pan-bone thinning of the lower jaws, mandibular fat pads, and isolation of the ear region), probably enabling them to hear the higher sonic frequencies of sound-producing fish on which they preyed. Ultrasonic echolocation evolved in Oligocene odontocetes, enabling them to find silent prey. Asymmetry and much of the sonic-frequency range of directional hearing were lost in Oligocene mysticetes during the shift to lowfrequency hearing and bulk-straining predation.Cetacea | land-to-sea transition M ost mammals have bilaterally symmetrical skulls. Symmetrical crania characterize the artiodactyls closely related to whales, and symmetrical crania characterize mysticetes within Cetacea (1) (Fig. 1A). Odontocetes are exceptional because most odontocete crania are asymmetrical, with dorsal cranial bones shifted posteriorly and to the left side (1-8). Living odontocetes have a hypertrophied melon, nasal sacs, and phonic lips used to produce high-frequency sound (> 20 kHz) (9-11). Mysticetes lack these specializations of the nasal apparatus, use low-frequency sound (11,12), and may use the larynx (13) to produce lowfrequency sound. Coupling of high-frequency echolocation with facial and cranial asymmetry in living odontocetes, and the absence of both in living artiodactyls and living mysticetes, make it reasonable to expect that asymmetry originated in odontocetes (5-7). However, it is unresolved how the cranial asymmetry of odontocetes evolved in the transition from archaeocetes to modern whales, and the history becomes even more complex when archaeocetes themselves are considered.Eocene archaeocete whales gave rise to all modern toothed and baleen whales during or near the Eocene-Oligocene transition (14-16). Archaeocetes were previously thought to have symmetrical skulls (3, 5, 7). Asymmetry observed in fossil crania has often been assumed to be an artifact of deformation following burial, and it has been ignored or even removed in published drawings [as was done initially for three of the skulls we studied (17-19)].
ResultsHere we document and quantify asymmetry in archaeocete crania. Further observations on exceptionally well-preserved archaeocete crania and dentaries suggest a link between cranial asymmetry and the ability to locate sound sources in water.We quantified midline suture deviation, δx, from a straight rostrocaudal axis [RC, after Ness...