Snake bioacoustics
Young
(2003)
reviews the ability of snakes to produce and detect sounds, while leaving
unresolved the existence of acoustical communication within any snake species. Relative to other vertebrates, snakes
have reduced ear morphology: the external ear is absent, the middle ear
contains a single ossicle, and the inner ear lacks
the round window. Vibrations in
the stapes (the sole remaining middle ear bone) create pressure waves in the perilymphatic fluid of the inner ear, and it is suggested
that high frequencies create an impedence-matching
problem in the middle ear; however, the function of this vibration transmission
system is incompletely understood.
Confounding understanding of snake auditory perception is the fact that
ear morphology shows no clear phylogenetic or
ecological pattern. Cochlear
amplification (increasing sensitivity and frequency range by altering
populations of hair cells) has not been investigated in snakes. Due to differences in the propagation
of sounds through different media, fossorial,
aquatic, and arboreal snakes are expected to display different ear structures;
no study has yet investigated the responses of snakes to aquatic pressure
waves.
Studies
of snake auditory perception indicate that sound waves are perceived directly;
these vibrations do not transmit to the substrate well because of the impedance
mismatch between air and ground.
Additionally, because medium absorption is very high for
substrate-propagated vibrations, detection distances for ground vibrations are
low. Auditory sensitivity for most
species occurs in a narrow range of low frequencies (200 to 400 Hz), although
snakes also display somatic hearing (the perception of vibrations from the body
surface rather than the inner ear), which is characterized by less amplitude
sensitivity but a greater frequency range. Interestingly, in a study from 1923, a researcher played
pure tones from a telephone to a group of rattlesnakes: ÒA momentary sounding of
the telephone would instantly start the rattling.Ó Due to the large latency and refractory periods of their
auditory neurons, snakes are poorly suited to make time-domain analyses of
vibrations or spatial localization of sound through temporal encoding.
Sound
production by snakes can occur by tail vibration (with or without a rattle), cloacal popping (compressing and releasing air out of the
anal opening, which may create a monopole), scale abrasion (with or without
specialized rasping scale projections), hissing (which generally shows no
evidence of temporal patterning, harmonics, or amplitude or frequency
modulation, although one species a laryngeal septum [Òvocal cordÓ]), or
growling (in which tracheal divirticula function as
resonators). In rattlesnakes, the
rattle functions as a dipole and produces a broadband sound from 2 to 20 kHz
(up to 50 kHz in one study). In
common with the sound-producing muscles of other vertebrates, the tail muscles
of rattlesnakes are very energetically efficient.
Young,
B. A. 2003. Snake bioacoustics: Toward a richer understanding of
the behavioral ecology of snakes. Quarterly Review of
Biology. 78:303-325.