When insects first took to land around 400 million years ago, they were presumably deaf (Gopfert & Hennig, 2015). Since then, hearing in insects has evolved independently no less
than 19 times throughout their evolutionary history. Only a relatively small number of insect species can hear, but within that group, “ear” form and location is highly varied (Yager, 1999).
Insects use their hearing to help them detect and avoid predators and for intraspecies communication, including the act of attracting a mate by acoustic signaling. Insects have hearing that is sensitive at high frequencies and over a broad frequency range, but these specifications are evolutionarily “tuned” to the range of frequencies that best allow the insect to hear their predator and communicate within their species. Most insects have two widely-spaced ears which aid the insect in determining the direction of sound. Additionally, some parasitoid insects listen to the singing of other insects to locate hosts for their parasitic offspring (Yager, 1999).
For evolutionary reasons to be discussed later, the location of hearing organs in insects is highly varied. Ears are often located on the legs, mesothorax, metathorax, various segments of the abdomen, wings, mouth parts, and, most commonly, the caudal thorax or rostral abdomen (Yager, 1999).
Two different types of ears exist within insects: the tympanal ear, a pressure receiver, and the antennal ear, a particle velocity detector.
Gopfert, Martin. Hennig, Matthias. Types of tympanal ears. Hearing in Insects, 09 August 2022
As a pressure receiver, the tympanal ear is effective over long distances. It consists of three basic parts: the tympanum, the tracheal sac, and the tympanal organ. The tympanum membrane, or eardrum, is a piece of very thin cuticle that responds to the pressure difference between its opposing sides (pressure of the external environment versus the pressure of the internal environment) by vibrating. The shape, size, thickness, and material of the tympanum affect the range of frequencies it mechanically responds to. The tracheal sac is an air-filled sac on the inner side of the tympanum. It is analogous to the middle ear of mammals. The contents of the tracheal sac need to have an acoustic impedance that matches that of the external medium, so it is filled with air within terrestrial insects. Lastly, the tympanal organ is what converts the mechanical signal of vibration into neural signals. It is attached to the tympanum by an attachment cell and is made up of a varying number of chordotonal sensilla. In insects, there is one bipolar neuron per chordotonal sensillum. The receptor potential in the bipolar neurons is stimulated by the vibrations of the tympanum. The axons of the bipolar neurons terminate in the central nervous system (CNS) in an auditory neuropil to synapse with interneurons (Yager, 1999). Hearing is thus achieved.
For this ear type, the antennae is vibrated by moving sound particles, not pressure waves. Thus, it is best used for close distances and low frequencies. Instead of the fluid-filled tracheal sac matching impedance, the antenna has a single resonant filter within. If frequencies get past this filter, they act as tuned input signals producing a neural response. Antennal ears have 2 or 3 bipolar neurons per chordotonal sensilla compared to the 1:1 of tympanal ears (Gopfert & Hennig, 2015).
The need for communication, a skill that first appeared around 165 million years ago (Gopfert & Hennig, 2015).
The need to evade predation by detecting predators. Most prominently, this refers to the detection of bats which use echolocation to locate their prey. Bat predation dates back to around 65 million years ago (Gopfert & Hennig, 2015).
The hearing organs of insects evolved from chordotonal organs, mechanosensory internal-stretch receptors. Chordotonal organs are responsible for monitoring the insects movement and can be found all along the body of the insect, explaining the wide variation in ear locations. Evolutionarily speaking, different species of insects “picked” a pair of chordotonal organs and transformed them into auditory organs by coupling them to structures that are able to vibrate in response to sound, like antennae or tympanum. This transformation also explains the multifunctionality of some insect species’ ears, as these organs sometimes have functions that extend past just hearing, given their mechanosensory ancestry. Currently, most insect species are still considered deaf. Others have lost their hearing abilities when they evolved into flightless insects and were no longer threatened by bat predation (Gopfert & Hennig, 2015).