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TitleEvolution of the Vertebrate Auditory System
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Page 2

SPRINGER HANDBOOK OF

AUDITORY RESEARCH

Series Editors: Richard R. Fay and Arthur N. Popper

Springer Science+ Business Media, LLC

Page 216

6. Evolution of the Amphibian Ear 199

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Page 217

7
The Lizard Basilar Papilla
and Its Evolution

GEOFFREY A. MANLEY

1. Introduction

Lizards are as structurally diverse as other groups of amniotes, such as mammals
and birds. When we look at their hearing organs, however, the structural variety
that we see in lizards exceeds that seen in any other amniote group. Its length
alone can vary up to a factor of more than 40 times. In fact, this may well be
the most variable sense organ seen in vertebrates (Manley 2000c,d).

In general, we associate structural variations with functional specializations.
For example, the auditory foveae [a region of the auditory papilla expanded to
accommodate a small but very important frequency region (Koppl 2001; Vater
2001)] of the barn owl and of some bat species clearly result from selection
pressures for passive or active sound localization. In almost all lizards, however,
there is no evidence for specialization to particular, known aspects of the species'
lifestyles, in spite of a family-, genus-, and even species-specific structure of the
hearing organ (Wever 1978; Miller 1980). This chapter shows that in spite of
this enormous structural variety, the hearing organ of lizards demonstrates sur-
prisingly little functional variation, which suggests that many of the structural
changes were selectively equivalent. The lizard ear is arguably an excellent case
to illustrate the controversial concept that has been termed "neutral" evolution,
i.e., in this case changes in structure that are not the result of any particular
selective pressure.

This chapter does not consider any changes that have occurred in the evolution
of the middle ear in different lizard families. A number of morphological
changes are observed in different groups, including loss of the tympanic
membrane. Wever (1978) identified three general types of lizard middle ear,
based partly on the structure of the extracolumella and the presence or absence
of tympanic muscles. Except for similarities in middle-ear structure between
related lizard families, however, few data provided insights into the evolution of
middle-ear diversity. Wever also studied the efficiency of sound transmission in
different species and found some substantial differences. Since then, little work
has been carried out on differences between lizard middle ears that is relevant
for asking questions about evolutionary trends.

200

Page 432

Type I, hair cells, 59-60
Type II, hair cells, 59
Type IV cell type, birds and mammals,

308
Tyto alba (barn owl), auditory midbrain

motor systems, 328-329
basilar papilla specializations, 70, 226,

232-233, 238ff
frequency map, 228ff
lID processing, 320
lTD processing, 313ff
nucleus larninaris, 312
octaval column cell types, 304ff
papillar length and hair cell number,

236ff
space-tuned neurons in forebrain, 338

Ultrasound, hearing in herrings, 99
Underwater hearing, amphibians, 164,

174
Unidirectional-type hair cells, reptiles,

169
Urochordates, ciliated mechanoreceptors,

77ff
cupular organs, 77-78

Urodeles, hair cells, 65ff

Index 415

Vertebrate, cladogram, 3
on geological time scale, 5
phylogeny, Iff
taxa hierarchy, 4

Vertebrate hair cells, summary, 73-74
Vestibular epithelia, 59ff
Vestibular labyrinth, evolution of hearing,

96ff
Violin, sound production, 34
Virginia opossum, see Didelphis

virginiana
Vocalization, and evolution of hearing,

328-329
archosaurs, 224ff
pathways in birds and mammals, 340-

341
sound processing in midbrain, 331-332

Weberian ossicles, 107ff, 128, 130
extirpation, 113

Xenopus laevis (African clawed frog),
ear, 173ff

auditory midbrain organization, 326
periotic canal, 179-181
underwater hearing, 174

Page 433

SPRINGER HANDBOOK OF AUDITORY RESEARCH (continued from page ii)

Volume 22: Evolution of the Vertebrate Auditory System
Edited by Geoffrey A. Manley, Arthur N. Popper and Richard R. Fay

For more information about the series, please visit www.springer-ny.comlshar.

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