Male anuran amphibians (frogs and toads) use auditory
(Bogert, 1960; Littlejohn, 1977; Wells, 1977; Gerhardt and
Schwartz, 1995) and visual (for a review, see Hödl and
Amézquita, 2001) cues to attract conspecific females and to
defend territories against potential conspecific intruders. In
most species of anuran, males are vocal and females are not,
representing a clear example of a behavioral difference
between the sexes. Males and females may also exhibit
external morphological and other differences. For example,
secondary sexual characteristics are traits other than those
associated with the gonads and their ducts that differ between
the sexes (Noble, 1931). The development of secondary sexual
characteristics, resulting in sexual dimorphism, is a common
phenomenon among the anurans (duToit, 1943). These
characteristics may persist throughout the adult life of the
animal; for example, the sexually dimorphic size of the
tympanic membrane in some anuran species, such as the North
American bullfrog Rana catesbeiana, in which the tympanic
membrane of the male is larger than that of the female (Shofner
and Feng, 1981; Hetherington, 1994; Purgue, 1997). Other
examples of permanent secondary sexual characteristics are the
bright coloration of males of Bufo periglenes compared with
the females (Savage, 1967) and the unique frontal swellings on
the head of male Rana pileata (Noble, 1931). In contrast, some
species have ephemeral secondary sexual characteristics that
appear only during the breeding season, in which case their
expression is presumably under hormonal control. These
include the nuptial pads of many male frogs and toads
(Klemens, 1997), the prepollical spines on males of Hyla
rosenbergi (Kluge, 1981) and the labial spines of
1223 The Journal of Experimental Biology 204, 1223–1232 (2001)
Printed in Great Britain © The Company of Biologists Limited 2001
JEB3235
During the breeding season, each tympanic membrane
of males of the Old World treefrog Petropedetes parkeri
is decorated with a single, prominent, fleshy tympanic
papilla. The tympanic papilla, located dorsally on the
tympanic membrane, is covered by an epidermal surface
and is composed of non-ossified, spongiform tissue
containing a number of globular, fluid-filled vesicles found
at highest density near the papillar tip. These vesicles
appear to have exit pores and are probably simple alveolar
exocrine glands. Injecting sound into the pressurized vocal
cavity of the male and measuring the vibration velocity
response of the tympanic membrane revealed that from 0.3
to 2.0 kHz the tympanic papilla velocity amplitude is on
average 20 dB lower than that of a point diametrically
opposite on the ventral half of the tympanic membrane.
The close agreement between the dominant frequency of
the call and the frequency of the maximum spectral peak
of the Fast Fourier Transform of the impulse response of
the eardrum is consistent with the use of the eardrum in
this species both as a call receiver and as a call radiator,
similar to the function suggested for the eardrum of the
male bullfrog Rana catesbeiana. Unexpectedly, surgically
removing the tympanic papilla lowered the frequency
of the peak vibrational amplitude, testifying to the
importance of membrane tension as a dominant factor in
the vibratory behavior of the eardrum. During normal
positive-pressure breathing, the tympanic papillae move
conspicuously, suggesting a possible rôle as a visual signal.
Key words: tympanic membrane, Ranidae, Cameroon, Petropedetes
parkeri, columella, tympanic papilla, communication.
Summary
Introduction
FUNCTIONAL CONSEQUENCES OF A NOVEL MIDDLE EAR ADAPTATION IN THE
CENTRAL AFRICAN FROG PETROPEDETES PARKERI (RANIDAE)
PETER M. NARINS
1,
*, EDWIN R. LEWIS
2
, ALEJANDRO P. PURGUE
1,
‡, PHILLIP J. BISHOP
3,
§,
LESLIE R. MINTER
4
AND DWIGHT P. LAWSON
5,
¶
1
Department of Physiological Science, University of California, Los Angeles, 405 Hilgard Avenue, Los Angeles,
CA 90095, USA,
2
Department of Electrical Engineering and Computer Science, University of California, Berkeley,
CA 94270, USA,
3
Department of Zoology, University of the Witwatersrand, Johannesburg, Republic of South Africa,
4
Department of Medical Science, University of the North, Sovenga, Republic of South Africa and
5
Department of
Biology, University of Texas at Arlington, Arlington, TX 76019, USA
*e-mail: pnarins@ucla.edu
‡Present address: Bioacoustics Research Program, 159 Sapsucker Woods Road, Ithaca, NY 14850, USA.
§Present address: Department of Zoology, University of Otago, Dunedin 9030, New Zealand.
¶Present address: Zoo Atlanta, 800 Cherokee Avenue S.E., Atlanta, GA 30315, USA.
Accepted 3 January; published on WWW 15 March 2001