The Origin of High-Frequency Hearing in Whales

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The Origin of High-Frequency Hearing in Whales Morgan Churchill, Manuel Martinez-Caceres, Christian de Muizon, Jessica Mnieckowski, Jonathan H. Geisler  Current Biology  Volume 26, Issue 16, Pages 2144-2149 (August 2016) DOI: 10.1016/j.cub.2016.06.004 Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 1 Holotype Skull, Mandible, and Right Bulla of Echovenator sandersi (A–N) Holotype mandible (A), atlas (B and C), cranium (D–K), and right tympanic bullae (L–N) of Echovenator sandersi. Cranium presented in dorsal (D and E), ventral (F and G), lateral (H and I), and anterior (J and K) views, with anatomical features labeled on the left and cranial bones indicated by color on the diagrams to the right. Abbreviations are as follows: afm, antorbital foramina of the maxilla; ang, angular process of the mandible; asl, ascending process of the lacrimal; asm, ascending process of the maxilla; con, occipital condyle; cp, coronoid process of the mandible; en, external nares; ep, embrasure pits; fps, falciform process of the squamosal; frf, frontal fossa; gf, glenoid fossa; ic, exposed incisivomaxillary canal; icn, intercondyloid notch; mf, mental foramina; mpc, medial prenarial crest; mr, mandibular ramus; ms, mandibular symphysis; nc, nuchal crest; npp, narial process of the premaxilla; paf, palatine foramen; pgp, postglenoid process; plf, posterior lacerate foramen (cranial hiatus); pnf, paranaris fossa; ppe, paraoccipital process of the exoccipital; prpf, preorbital process of the frontal; ps; posterior sinus; pspf, postorbital process of the frontal; ptc, parasagittal temporal crest; sopf, supraorbital process of the frontal; sqf, squamosal fossa; stc, subtemporal crest; sul, supraorbital sulcus of the maxilla; VIII, exit of the mandibular branch of the trigeminal nerve VIII; zyg, zygomatic process of the squamosal. Colors used to indicate different bones are as follows: cream, frontal; red, nasals; teal green, parietals; light purple, alisphenoid; light blue, basioccipital; blue green, mandible; dark blue, exoccipital and supraoccipital; dark orange, jugal; orange, maxilla; yellow, palatine; dark green, lacrimal; light green, premaxilla; dark pink, presphenoid and basisphenoid; dark purple, squamosal; dark red, vomer; brown, basioccipital. Dark gray-shaded regions indicate reconstructed parts with patching putty. Dashed lines and hatched regions indicate missing parts of the specimen. See also Supplemental Experimental Procedures, Figures S1 and S2, and Table S1. Current Biology 2016 26, 2144-2149DOI: (10.1016/j.cub.2016.06.004) Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 2 Right Periotic of Echovenator sandersi (A–F) Right periotic of Echovenator sandersi in ventrolateral (A and D), dorsal (B and E) and medial (C and F) views. Abbreviations are as follows: aes, anteroexternal sulcus; app, anterior process of the periotic; cc, cochlear crest (posterolateral crest of Basilosauridae); cn, foramen for cochlear nerve (branch of the vestibulo-cochlear nerve, VIII) and tractus spiralis foraminosus; dc, dorsal crest (superior process or tegmen tympani); elf, foramen for the endolymphatic canal; eth, epitympanic hiatus; fc, fenestra cochleae (round window); fg, groove for the facial nerve; fi, fossa incudi; fn, internal foramen for the facial nerve (VII); fm, fossa for the malleus head; fst, fossa for the stapedial muscle; gtt, groove for the tensor tympani muscle; iam, internal auditory meatus; plf, foramen for the cochlear aqueduct (perilymphatic canal); ppp, posterior process of the periotic; pr, promontorium; pyr, pyramidal process; smf, suprameatal fossa; st, stapes; vlt, ventrolateral tuberosity of the periotic; vn, foramen for the vestibular nerve (branch of the vestibulo-cochlear nerve, VIII). Internal morphology of the inner ear shown in Figure S1. Current Biology 2016 26, 2144-2149DOI: (10.1016/j.cub.2016.06.004) Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 3 Simplified Phylogeny Indicating the Phylogenetic Relationships for Echovenator sandersi, Showing the Evolution of Hearing-Related Traits Whales represented in color text, with archaeocetes in green, mysticetes in red, and odontocetes in blue, with Echovenator highlighted in bold. Horizontal bars near inner ears are scale bars (5 mm), with hearing ranges indicated in red text, while black bars and text indicate the origin of specific inner ear features based on ancestral character state reconstruction. Dashed lines for characters represent regions of the tree where the ancestral character state reconstruction is ambiguous for a given trait. Approximate maximum apical extent of SLA from base of the first cochlear turn delimited by yellow arc, with an arrow indicating this feature in Piscobalaena. CL refers to cochlear length, while FC refers to area of the fenestra cochlearis. Isosurface renderings of inner ears presented in apical (top) and medial (middle) views, with the cochlea shaded blue and other inner ear structures shaded red. Piscobalaena and Phocoena are represented by left ears that have been flipped to ease comparison with the other renderings. Complete phylogeny is presented in Figure S3, and character matrix is included within the supplemental material. Current Biology 2016 26, 2144-2149DOI: (10.1016/j.cub.2016.06.004) Copyright © 2016 Elsevier Ltd Terms and Conditions

Figure 4 PCA of Nine Measurements of the Inner Ear of Whales and Their Close Relatives PC 1 represents variation in body size, while PC 2 represents variation in morphological features associated with high- and low-frequency hearing. Ultrasonic hearing morphospace is indicated in light gray and infrasonic in dark gray. Symbols represent hippopotamids (white squares), archaeocetes (black triangles), fossil mysticetes (black diamonds), extant mysticetes (white diamonds), fossil odontocetes (black circles), extant odontocetes (white circles), and Echovenator sandersi (star). See also Figure S4 and Table S2. Current Biology 2016 26, 2144-2149DOI: (10.1016/j.cub.2016.06.004) Copyright © 2016 Elsevier Ltd Terms and Conditions