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@ -170,7 +170,8 @@
volume={122}, volume={122},
pages={53--64}, pages={53--64},
year={1977}, year={1977},
} }# Cited
@article{helversen1984parallel, @article{helversen1984parallel,
title={{Parallel processing in auditory pattern recognition and directional analysis by the grasshopper Chorthippus biguttulus L. (Acrididae)}}, title={{Parallel processing in auditory pattern recognition and directional analysis by the grasshopper Chorthippus biguttulus L. (Acrididae)}},
author={von Helversen, Dagmar}, author={von Helversen, Dagmar},
@ -207,7 +208,8 @@
pages={373--386}, pages={373--386},
year={1997}, year={1997},
publisher={Springer} publisher={Springer}
} }# Cited
@article{helversen2004acoustic, @article{helversen2004acoustic,
title={{Acoustic communication in a duetting grasshopper: Receiver response variability, male strategies and signal design}}, title={{Acoustic communication in a duetting grasshopper: Receiver response variability, male strategies and signal design}},
author={von Helversen, Dagmar and Balakrishnan, Rohini and von Helversen, Otto}, author={von Helversen, Dagmar and Balakrishnan, Rohini and von Helversen, Otto},
@ -365,7 +367,8 @@
author={Otte, Daniel}, author={Otte, Daniel},
year={1970}, year={1970},
publisher={University of Michigan Museum of Zoology} publisher={University of Michigan Museum of Zoology}
} }# Cited
@article{prinz2002temporal, @article{prinz2002temporal,
title={Temporal modulation transfer functions in auditory receptor fibres of the locust (Locusta migratoria L.)}, title={Temporal modulation transfer functions in auditory receptor fibres of the locust (Locusta migratoria L.)},
author={Prinz, P and Ronacher, B}, author={Prinz, P and Ronacher, B},

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@ -7,25 +7,58 @@
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118
main.bbl
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@ -403,12 +403,12 @@
\endentry \endentry
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@ -2365,29 +2365,39 @@
<bcf:citekey order="1" intorder="1">helversen1972gesang</bcf:citekey> <bcf:citekey order="1" intorder="1">helversen1972gesang</bcf:citekey>
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<bcf:citekey order="4" intorder="1">vedenina2011speciation</bcf:citekey> <bcf:citekey order="4" intorder="1">otte1970comparative</bcf:citekey>
<bcf:citekey order="5" intorder="1">sevastianov2023evolution</bcf:citekey> <bcf:citekey order="5" intorder="1">vedenina2011speciation</bcf:citekey>
<bcf:citekey order="6" intorder="1">cigliano2018orthoptera</bcf:citekey> <bcf:citekey order="6" intorder="1">sevastianov2023evolution</bcf:citekey>
<bcf:citekey order="7" intorder="1">clemens2013computational</bcf:citekey> <bcf:citekey order="7" intorder="1">helversen1977stridulatory</bcf:citekey>
<bcf:citekey order="8" intorder="1">clemens2013feature</bcf:citekey> <bcf:citekey order="8" intorder="1">helversen1997recognition</bcf:citekey>
<bcf:citekey order="9" intorder="1">ronacher2015computational</bcf:citekey> <bcf:citekey order="9" intorder="1">otte1970comparative</bcf:citekey>
<bcf:citekey order="10" intorder="1">rehbein1974structure</bcf:citekey> <bcf:citekey order="10" intorder="1">helversen1977stridulatory</bcf:citekey>
<bcf:citekey order="11" intorder="1">kalmring1975afferent</bcf:citekey> <bcf:citekey order="11" intorder="1">helversen1997recognition</bcf:citekey>
<bcf:citekey order="12" intorder="1">rehbein1976auditory</bcf:citekey> <bcf:citekey order="12" intorder="1">vedenina2011speciation</bcf:citekey>
<bcf:citekey order="13" intorder="1">eichendorf1980projections</bcf:citekey> <bcf:citekey order="13" intorder="1">sevastianov2023evolution</bcf:citekey>
<bcf:citekey order="14" intorder="1">clemens2011efficient</bcf:citekey> <bcf:citekey order="14" intorder="1">helversen1977stridulatory</bcf:citekey>
<bcf:citekey order="15" intorder="1">clemens2011efficient</bcf:citekey> <bcf:citekey order="15" intorder="1">helversen1997recognition</bcf:citekey>
<bcf:citekey order="16" intorder="1">clemens2011efficient</bcf:citekey> <bcf:citekey order="16" intorder="1">cigliano2018orthoptera</bcf:citekey>
<bcf:citekey order="17" intorder="1">michelsen1971frequency</bcf:citekey> <bcf:citekey order="17" intorder="1">clemens2013computational</bcf:citekey>
<bcf:citekey order="18" intorder="1">windmill2008time</bcf:citekey> <bcf:citekey order="18" intorder="1">clemens2013feature</bcf:citekey>
<bcf:citekey order="19" intorder="1">malkin2014energy</bcf:citekey> <bcf:citekey order="19" intorder="1">ronacher2015computational</bcf:citekey>
<bcf:citekey order="20" intorder="1">machens2001discrimination</bcf:citekey> <bcf:citekey order="20" intorder="1">rehbein1974structure</bcf:citekey>
<bcf:citekey order="21" intorder="1">machens2001representation</bcf:citekey> <bcf:citekey order="21" intorder="1">kalmring1975afferent</bcf:citekey>
<bcf:citekey order="22" intorder="1">suga1960peripheral</bcf:citekey> <bcf:citekey order="22" intorder="1">rehbein1976auditory</bcf:citekey>
<bcf:citekey order="23" intorder="1">gollisch2002energy</bcf:citekey> <bcf:citekey order="23" intorder="1">eichendorf1980projections</bcf:citekey>
<bcf:citekey order="24" intorder="1">hildebrandt2009origin</bcf:citekey> <bcf:citekey order="24" intorder="1">clemens2011efficient</bcf:citekey>
<bcf:citekey order="25" intorder="1">clemens2010intensity</bcf:citekey> <bcf:citekey order="25" intorder="1">clemens2011efficient</bcf:citekey>
<bcf:citekey order="26" intorder="1">fisch2012channel</bcf:citekey> <bcf:citekey order="26" intorder="1">clemens2011efficient</bcf:citekey>
<bcf:citekey order="27" intorder="1">michelsen1971frequency</bcf:citekey>
<bcf:citekey order="28" intorder="1">windmill2008time</bcf:citekey>
<bcf:citekey order="29" intorder="1">malkin2014energy</bcf:citekey>
<bcf:citekey order="30" intorder="1">machens2001discrimination</bcf:citekey>
<bcf:citekey order="31" intorder="1">machens2001representation</bcf:citekey>
<bcf:citekey order="32" intorder="1">suga1960peripheral</bcf:citekey>
<bcf:citekey order="33" intorder="1">gollisch2002energy</bcf:citekey>
<bcf:citekey order="34" intorder="1">hildebrandt2009origin</bcf:citekey>
<bcf:citekey order="35" intorder="1">clemens2010intensity</bcf:citekey>
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@ -81,60 +81,61 @@ distributed accumulation of anatomical, physiological and ethological evidence.
This process is undoubtedly without alternative; however, it leaves us with the This process is undoubtedly without alternative; however, it leaves us with the
challenge of integrating the available fragments into a coherent whole in order challenge of integrating the available fragments into a coherent whole in order
to address issues such as the interaction between individual system components, to address issues such as the interaction between individual system components,
the functional limitations of the system overall, or taxonomic comparisons of the functional limitations of the system overall, or taxonomic comparisons
systems that process the same sensory modality. Any unified framework that between systems that process the same sensory modality. Any unified framework
captures the essential functional aspects of a given sensory system thus has that captures the essential functional aspects of a given sensory system thus
the potential to deepen our current understanding and fasciliate systematic has the potential to deepen our current understanding and fasciliate systematic
investigations. However, building such a framework is a challenging task. It investigations. However, building such a framework is a challenging task. It
requires a wealth of existing knowledge of the system and the signals it requires a wealth of existing knowledge of the system and the signals it
operates on, a clearly defined scope, and careful reduction, abstraction, and operates on, a clearly defined scope, and careful reduction, abstraction, and
formalization of the underlying anatomical structures and physiological formalization of the underlying structures and mechanisms.
mechanisms.
One sensory system about which extensive information has been gathered over the One sensory system about which extensive information has been gathered over the
years is the auditory system of grasshoppers~(\textit{Acrididae}). Grasshoppers years is the auditory system of grasshoppers~(\textit{Acrididae}). Grasshoppers
rely on auditory processing primarily for intraspecific communication, which rely on their sense of hearing primarily for intraspecific communication, which
includes mate attraction and evaluation~(\bcite{helversen1972gesang}), sender includes mate attraction and evaluation~(\bcite{helversen1972gesang}), sender
localization~(\bcite{helversen1988interaural}), courtship display~(SOURCE), localization~(\bcite{helversen1988interaural}), courtship display~(SOURCE),
rival deterrence~(\bcite{greenfield1993acoustic}), and loss-of-signal predator rival deterrence~(\bcite{greenfield1993acoustic}), and loss-of-signal predator
alarm~(SOURCE). The different behavioral contexts are met with alarm~(SOURCE). In accordance with this rich behavioral repertoire,
grasshoppers have evolved a variety of sound production mechanisms to generate
acoustic communication signals for different contexts and ranges using their
wings, hindlegs, or mandibles~(\bcite{otte1970comparative}). Among the most
conspicuous acoustic signals of grasshoppers are the species-specific calling
songs, which broadcast the presence of the singing individual --- usually the
males of a species --- to potential mates within range. These songs are usually
more characteristic of a species than morphological traits.
Different acustic signals are used for different behavioral This reliance on species-specific acoustic signals for mediating
contexts and communication ranges reproduction represents a strong evolutionary driving force, that resulted in a
massive species diversification~(\bcite{vedenina2011speciation},
\bcite{sevastianov2023evolution}).
Depending on the behavioral context and the communication range, The characteristic calling songs are
produced by stridulation, during which the grasshopper pulls the serrated
stridulatory file on its hindlegs across a resonating vein on the
forewings~(\bcite{helversen1977stridulatory},
\bcite{helversen1997recognition}).
Such elaborate acoustic behaviors co-depend not only on
Grasshoppers generate their most conspicious acoustic signals reliable auditory perception but also on suitable acoustic signals for
---~commonly referred to as "songs"~--- by stridulation. different contexts and ranges. To this end, grasshoppers have evolved a variety
of sound production mechanisms using their wings, hindlegs, or
mandibles~(\bcite{otte1970comparative}). The most conspicuous acoustic signals
Different acoustic signals may be generated using different --- the characteristic calling songs --- are produced by stridulation, during
body parts ---~wings, hindlegs, or mandibles~--- which the grasshopper pulls the serrated stridulatory file on its hindlegs
across a resonating vein on the forewings~(\bcite{helversen1977stridulatory},
\bcite{helversen1997recognition}). The reliance on species-specific acoustic
Different acoustic signals may be generated using different communication signals represents a strong evolutionary driving force, that
body parts ---~wings, hindlegs, or mandibles~--- but the most conspicious resulted in a massive species diversification~(\bcite{vedenina2011speciation},
The required acoustic signals for different contexts and ranges
may be generated using different body parts ---~wings, hindlegs, or
mandibles~--- but the most common sound production mechanism is stridulation,
during which the animal pulls the serrated stridulatory file on its hindlegs
across a resonating vein on the forewings. The resulting "songs"
The reliance on acoustic communication signals represents a strong evolutionary
driving force, that resulted in a massive species diversification among
grasshoppers~(\bcite{vedenina2011speciation},
\bcite{sevastianov2023evolution}). \bcite{sevastianov2023evolution}).
The most conspicuous acoustic signals
Grasshoppers produce their most conspicious acoustic signals --- the characteristic calling songs --- are produced by stridulation, during
---~commonly referred to as "songs"~--- by stridulation, during which the which the grasshopper pulls the serrated stridulatory file on its hindlegs
animal rubs the serrated stridulatory file on its hindleg across a resonating across a resonating vein on the forewings~(\bcite{helversen1977stridulatory},
vein on the forewing. \bcite{helversen1997recognition}). Grasshopper songs are species-specific,
amplitude-modulated (AM) broad-band acoustic signals
Among the several thousand recognized grasshopper Among the several thousand recognized grasshopper
species~(\bcite{cigliano2018orthoptera}), diverse species-specific sound species~(\bcite{cigliano2018orthoptera}), diverse species-specific sound
@ -148,6 +149,10 @@ Strong dependence on acoustic signals for ranged communication\\
- Mate attraction/evaluation, rival deterrence, loss-of-signal predator alarm\\ - Mate attraction/evaluation, rival deterrence, loss-of-signal predator alarm\\
$\rightarrow$ Elaborate acoustic behaviors co-depend on reliable auditory perception $\rightarrow$ Elaborate acoustic behaviors co-depend on reliable auditory perception
Songs = Amplitude-modulated (AM) broad-band acoustic signals\\ Songs = Amplitude-modulated (AM) broad-band acoustic signals\\
- Generated by stridulatory movement of hindlegs against forewings\\ - Generated by stridulatory movement of hindlegs against forewings\\
- Shorter time scales: Characteristic temporal waveform pattern\\ - Shorter time scales: Characteristic temporal waveform pattern\\