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@ -39,10 +39,11 @@
pages={619--629},
year={1985},
}
@misc{cigliano2018orthoptera,
title={{Orthoptera Species File. Version 5.0/5.0}},
author={Cigliano, MM and Braun, H and Eades, DC and Otte, D},
year={2018}
@article{cigliano2024orthoptera,
title={Orthoptera species file},
author={Cigliano, Mar{\'\i}a Marta and Braun, Holger and Eades, David C and Otte, Daniel},
year={2024},
publisher={Species File Group}
}# Cited
@article{clemens2013computational,
@ -179,7 +180,8 @@
volume={154},
pages={837--846},
year={1984},
}
}# Cited
@article{helversen1988interaural,
title={Interaural intensity and time discrimination in an unrestraint grasshopper: a tentative behavioural approach},
author={von Helversen, Dagmar and Rheinlaender, J{\"u}rgen},
@ -198,7 +200,8 @@
volume={172},
pages={633--639},
year={1993},
}
}# Cited
@article{helversen1997recognition,
title={Recognition of sex in the acoustic communication of the grasshopper Chorthippus biguttulus (Orthoptera, Acrididae)},
author={von Helversen, Dagmar and von Helversen, Otto},
@ -283,6 +286,14 @@
year={1978},
publisher={Springer}
}
@article{kohler2017morphological,
title={{Morphological and colour morph clines along an altitudinal gradient in the meadow grasshopper Pseudochorthippus parallelus}},
author={K{\"o}hler, G{\"u}nter and Samietz, J{\"o}rg and Schielzeth, Holger},
journal={PLoS ONE},
volume={12},
pages={e0189815},
year={2017},
}# Cited
@article{lang2000acoustic,
title={Acoustic communication distances of a gomphocerine grasshopper},
@ -354,6 +365,26 @@
year={1978},
publisher={Springer}
}
@article{miller1977directional,
title={Directional hearing in the locust Schistocerca gregaria Forsk{\aa}l (Acrididae, Orthophera)},
author={Miller, Lee A},
journal={Journal of comparative physiology},
volume={119},
number={1},
pages={85--98},
year={1977},
publisher={Springer}
}
@article{minckley1995psychoacoustics,
title={Psychoacoustics of female phonotaxis and the evolution of male signal interactions in Orthoptera},
author={Minckley, RL and Greenfield, MD},
journal={Ethology ecology \& evolution},
volume={7},
number={3},
pages={235--243},
year={1995},
publisher={Taylor \& Francis}
}
@article{neuhofer2008evolutionarily,
title={Evolutionarily conserved coding properties of auditory neurons across grasshopper species},
author={Neuhofer, Daniela and Wohlgemuth, Sandra and Stumpner, Andreas and Ronacher, Bernhard},
@ -428,7 +459,8 @@
volume={158},
pages={363--374},
year={1986},
}
}# Cited
@article{ronacher2008discrimination,
title={Discrimination of acoustic communication signals by grasshoppers (Chorthippus biguttulus): temporal resolution, temporal integration, and the impact of intrinsic noise.},
author={Ronacher, Bernhard and Wohlgemuth, Sandra and Vogel, Astrid and Krahe, R{\"u}diger},
@ -457,6 +489,16 @@
pages={61--71},
year={2015},
}
@incollection{rowell1972variable,
title={The variable coloration of the acridoid grasshoppers.},
author={Rowell, CH Fraser},
booktitle={Advances in Insect Physiology},
volume={8},
pages={145--198},
year={1972},
publisher = {Academic Press},
}# Cited
@article{sevastianov2023evolution,
title={{Evolution of calling songs in the grasshopper subfamily Gomphocerinae (Orthoptera, Acrididae)}},
author={Sevastianov, Nikita and Neretina, Tatiana and Vedenina, Varvara},
@ -464,7 +506,8 @@
volume={52},
pages={154--175},
year={2023},
}
}# Cited
@article{sippel1983nonlinear,
title={Non-linear analysis of the transmission of signals in the auditory system of the migratory locust Locusta migratorta},
author={Sippel, Martin and Breckow, Joachim},
@ -492,7 +535,8 @@
volume={6},
pages={1--23},
year={1994},
}
}# Cited
@article{suga1960peripheral,
title={Peripheral mechanism of hearing in locust},
author={Suga, Nobuo},
@ -517,7 +561,8 @@
volume={4965},
pages={244--260},
year={2021},
}
}# Cited
@article{tishechkin2016acoustic,
title={Acoustic signals in insects: A reproductive barrier and a taxonomic character},
author={Tishechkin, D Yu and Vedenina, V Yu},
@ -525,7 +570,8 @@
volume={96},
pages={1127--1164},
year={2016},
}
}# Cited
@article{vedenina2014stable,
title={{Stable and variable parameters in courtship songs of grasshoppers of the subfamily Gomphocerinae (Orthoptera, Acrididae)}},
author={Vedenina, V Yu and Shestakov, LS},
@ -549,7 +595,8 @@
volume={20},
pages={109--125},
year={2011},
}
}# Cited
@article{vedenina2013narrow,
title={{A narrow hybrid zone between the grasshoppers Stenobothrus clavatus and Stenobothrus rubicundus (Orthoptera: Gomphocerinae): Female preferences for courtship songs}},
author={Vedenina, Varvara Yu and F{\"a}hsing, Sylvia and Sradnick, Jan and Kl{\"o}pfel, Anja and Elsner, Norbert},

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@ -19,47 +19,50 @@
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<bcf:citekey order="2" intorder="1">helversen1988interaural</bcf:citekey>
<bcf:citekey order="3" intorder="1">greenfield1993acoustic</bcf:citekey>
<bcf:citekey order="4" intorder="1">otte1970comparative</bcf:citekey>
<bcf:citekey order="5" intorder="1">vedenina2011speciation</bcf:citekey>
<bcf:citekey order="6" intorder="1">sevastianov2023evolution</bcf:citekey>
<bcf:citekey order="7" intorder="1">helversen1977stridulatory</bcf:citekey>
<bcf:citekey order="8" intorder="1">helversen1997recognition</bcf:citekey>
<bcf:citekey order="9" intorder="1">otte1970comparative</bcf:citekey>
<bcf:citekey order="10" intorder="1">helversen1977stridulatory</bcf:citekey>
<bcf:citekey order="11" intorder="1">helversen1997recognition</bcf:citekey>
<bcf:citekey order="12" intorder="1">vedenina2011speciation</bcf:citekey>
<bcf:citekey order="13" intorder="1">sevastianov2023evolution</bcf:citekey>
<bcf:citekey order="14" intorder="1">helversen1977stridulatory</bcf:citekey>
<bcf:citekey order="15" intorder="1">helversen1997recognition</bcf:citekey>
<bcf:citekey order="16" intorder="1">cigliano2018orthoptera</bcf:citekey>
<bcf:citekey order="17" intorder="1">clemens2013computational</bcf:citekey>
<bcf:citekey order="18" intorder="1">clemens2013feature</bcf:citekey>
<bcf:citekey order="19" intorder="1">ronacher2015computational</bcf:citekey>
<bcf:citekey order="20" intorder="1">rehbein1974structure</bcf:citekey>
<bcf:citekey order="21" intorder="1">kalmring1975afferent</bcf:citekey>
<bcf:citekey order="22" intorder="1">rehbein1976auditory</bcf:citekey>
<bcf:citekey order="23" intorder="1">eichendorf1980projections</bcf:citekey>
<bcf:citekey order="24" intorder="1">clemens2011efficient</bcf:citekey>
<bcf:citekey order="25" intorder="1">clemens2011efficient</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>
<bcf:citekey order="36" intorder="1">fisch2012channel</bcf:citekey>
<bcf:citekey order="2" intorder="1">helversen1993absolute</bcf:citekey>
<bcf:citekey order="3" intorder="1">helversen1997recognition</bcf:citekey>
<bcf:citekey order="4" intorder="1">helversen1988interaural</bcf:citekey>
<bcf:citekey order="5" intorder="1">greenfield1993acoustic</bcf:citekey>
<bcf:citekey order="6" intorder="1">otte1970comparative</bcf:citekey>
<bcf:citekey order="7" intorder="1">tishechkin2016acoustic</bcf:citekey>
<bcf:citekey order="8" intorder="1">tarasova2021eurasius</bcf:citekey>
<bcf:citekey order="9" intorder="1">rowell1972variable</bcf:citekey>
<bcf:citekey order="10" intorder="1">kohler2017morphological</bcf:citekey>
<bcf:citekey order="11" intorder="1">vedenina2011speciation</bcf:citekey>
<bcf:citekey order="12" intorder="1">sevastianov2023evolution</bcf:citekey>
<bcf:citekey order="13" intorder="1">cigliano2024orthoptera</bcf:citekey>
<bcf:citekey order="14" intorder="1">helversen1984parallel</bcf:citekey>
<bcf:citekey order="15" intorder="1">ronacher1986routes</bcf:citekey>
<bcf:citekey order="16" intorder="1">helversen1988interaural</bcf:citekey>
<bcf:citekey order="17" intorder="1">helversen1977stridulatory</bcf:citekey>
<bcf:citekey order="18" intorder="1">stumpner1994song</bcf:citekey>
<bcf:citekey order="19" intorder="1">helversen1997recognition</bcf:citekey>
<bcf:citekey order="20" intorder="1">clemens2013computational</bcf:citekey>
<bcf:citekey order="21" intorder="1">clemens2013feature</bcf:citekey>
<bcf:citekey order="22" intorder="1">ronacher2015computational</bcf:citekey>
<bcf:citekey order="23" intorder="1">rehbein1974structure</bcf:citekey>
<bcf:citekey order="24" intorder="1">kalmring1975afferent</bcf:citekey>
<bcf:citekey order="25" intorder="1">rehbein1976auditory</bcf:citekey>
<bcf:citekey order="26" intorder="1">eichendorf1980projections</bcf:citekey>
<bcf:citekey order="27" intorder="1">clemens2011efficient</bcf:citekey>
<bcf:citekey order="28" intorder="1">clemens2011efficient</bcf:citekey>
<bcf:citekey order="29" intorder="1">clemens2011efficient</bcf:citekey>
<bcf:citekey order="30" intorder="1">michelsen1971frequency</bcf:citekey>
<bcf:citekey order="31" intorder="1">windmill2008time</bcf:citekey>
<bcf:citekey order="32" intorder="1">malkin2014energy</bcf:citekey>
<bcf:citekey order="33" intorder="1">machens2001discrimination</bcf:citekey>
<bcf:citekey order="34" intorder="1">machens2001representation</bcf:citekey>
<bcf:citekey order="35" intorder="1">suga1960peripheral</bcf:citekey>
<bcf:citekey order="36" intorder="1">gollisch2002energy</bcf:citekey>
<bcf:citekey order="37" intorder="1">hildebrandt2009origin</bcf:citekey>
<bcf:citekey order="38" intorder="1">clemens2010intensity</bcf:citekey>
<bcf:citekey order="39" intorder="1">fisch2012channel</bcf:citekey>
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@ -93,55 +93,40 @@ formalization of the underlying structures and mechanisms.
One sensory system about which extensive information has been gathered over the
years is the auditory system of grasshoppers~(\textit{Acrididae}). Grasshoppers
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}, \bcite{helversen1993absolute},
\bcite{helversen1997recognition}), sender
localization~(\bcite{helversen1988interaural}), courtship display~(SOURCE),
rival deterrence~(\bcite{greenfield1993acoustic}), and loss-of-signal predator
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.
This reliance on species-specific acoustic signals for mediating
conspicuous acoustic signals of grasshoppers are their species-specific calling
songs, which broadcast the presence of the singing individual --- mostly the
males of the species --- to potential mates within range. These songs are
usually more characteristic of a species than morphological
traits~(\bcite{tishechkin2016acoustic}, \bcite{tarasova2021eurasius}), which
can vary greatly within species~(\bcite{rowell1972variable},
\bcite{kohler2017morphological}). The reliance on acoustic signals to mediate
reproduction represents a strong evolutionary driving force, that resulted in a
massive species diversification~(\bcite{vedenina2011speciation},
\bcite{sevastianov2023evolution}).
\bcite{sevastianov2023evolution}), with over 6800 recognized grasshopper
species in the \textit{Acrididae} family~(\bcite{cigliano2024orthoptera}). It
is this diversity of species, and the crucial role of acoustic communication in
its emergence, that makes the grasshopper auditory system an intriguing
candidate for attempting to construct a functional model framework. For
simplicity, we focus on the pathway responsible for the recognition of
species-specific calling songs, disregarding other auditory functions such as
directional hearing~(\bcite{helversen1984parallel}, \bcite{ronacher1986routes},
\bcite{helversen1988interaural}).
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},
forewings~(\bcite{helversen1977stridulatory}, \bcite{stumpner1994song},
\bcite{helversen1997recognition}).
Such elaborate acoustic behaviors co-depend not only on
reliable auditory perception but also on suitable acoustic signals for
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
--- 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}). The reliance on species-specific acoustic
communication signals represents a strong evolutionary driving force, that
resulted in a massive species diversification~(\bcite{vedenina2011speciation},
\bcite{sevastianov2023evolution}).
The most conspicuous acoustic signals
--- 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}). Grasshopper songs are species-specific,
amplitude-modulated (AM) broad-band acoustic signals
Among the several thousand recognized grasshopper
species~(\bcite{cigliano2018orthoptera}), diverse species-specific sound
repertoires and production mechanisms
Strong dependence on acoustic signals for ranged communication\\
- Diverse species-specific sound repertoires and production mechanisms\\
@ -149,10 +134,6 @@ Strong dependence on acoustic signals for ranged communication\\
- Mate attraction/evaluation, rival deterrence, loss-of-signal predator alarm\\
$\rightarrow$ Elaborate acoustic behaviors co-depend on reliable auditory perception
Songs = Amplitude-modulated (AM) broad-band acoustic signals\\
- Generated by stridulatory movement of hindlegs against forewings\\
- Shorter time scales: Characteristic temporal waveform pattern\\