01878nas a2200253 4500008004100000022001400041245014500055210006900200260001600269300001200285490000600297520105900303653001701362653001701379653001901396653001101415653001401426653001501440100003301455700002201488700002401510700002201534856006801556 2011 eng d a1477-200000aQuality calls: phylogeny and biogeography of a new genus of neotropical katydid (Orthoptera: Tettigoniidae) with ultra pure-tone ultrasonics0 aQuality calls phylogeny and biogeography of a new genus of neotr cSep-03-2012 a77 - 940 v93 a

This paper describes Artiotonus, a new genus of tropical katydid from Colombia and Ecuador. These acoustic ensiferans are represented by three species with a geographic distribution generally restricted to the rainforest of the Bolivar geosyncline of northwestern South America (Pacific). A phylogenetic analysis based on 28 morphological and six behavioural characters produced a tree (A. artius, (A. tinae, A. captivus)) with a consistency index of 0.9. All species are defined by a few autapomorphic changes. The most reliable character for identification is the temporal pattern of the calling song, suggesting a recent genetic divergence. Biogeographic analysis indicates that such genetic divergence began with geographic isolation produced before the Holocene transgression. Species of this genus are remarkable in that the calling song of males exhibits a narrow spectrum with a high quality factor (37–42), unusual values for such ultrasonic signals. A broad discussion on the evolution of tonal calls and pulse trains is offered.

10aBiogeography10acalling song10aConocephalinae10ahigh-Q10apure tone10aultrasound1 aMontealegre-Zapata, Fernando1 aMorris, Glenn, K.1 aSarria-S, Fabio, A.1 aMason, Andrew, C. uhttp://www.tandfonline.com/doi/abs/10.1080/14772000.2011.56020902113nas a2200145 4500008004100000245013500041210006900176260001600245520156700261100003301828700001901861700002001880700002401900856004301924 2017 eng d00aMorphological determinants of carrier frequency signal in katydids (Orthoptera): a comparative analysis using biophysical evidence0 aMorphological determinants of carrier frequency signal in katydi cApr-09-20183 a

Male katydids produce mating calls by stridulation using specialized structures on the forewings. The right wing (RW) bears a scraper connected to a drum-like cell known as the mirror and a left wing (LW) that overlaps the RW and bears a serrated vein on the ventral side, the stridulatory file. Sound is generated with the scraper sweeping across the file, producing vibrations that are amplified by the mirror. Using this sound generator, katydids exploit a range of song carrier frequencies (CF) unsurpassed by any other insect group, with species singing as low as 600 Hz and others as high as 150 kHz. Sound generator size has been shown to scale negatively with CF, but such observations derive from studies based on few species, without phylogenetic control, and/or using only the RW mirror length. We carried out a phylogenetic comparative analysis involving 94 species of katydids to study the relationship between LW and RW components of the sound generator and the CF of the male's mating call, while taking into account body size and phylogenetic relationships. The results showed that CF negatively scaled with all morphological measures, but was most strongly related to components of the sound generation system (file, LW and RW mirrors). Interestingly, the LW mirror (reduced and non-functional) predicted CF more accurately than the RW mirror, and body size is not a reliable CF predictor. Mathematical models were verified on known species for predicting CF in species for which sound is unknown (e.g. fossils or museum specimens).

1 aMontealegre-Zapata, Fernando1 aOgden, Jessica1 aJonsson, Thorin1 aSoulsbury, Carl, D. uhttp://doi.wiley.com/10.1111/jeb.1317902082nas a2200229 4500008004100000022001400041245007400055210006900129260001600198300001600214490000700230520136400237653002301601653001601624653003101640653002001671100002901691700002501720700003301745700001901778856005501797 2011 eng d a0028-104200aTympanal mechanics and neural responses in the ears of a noctuid moth0 aTympanal mechanics and neural responses in the ears of a noctuid cJan-12-2011 a1057 - 10610 v983 a

Ears evolved in many groups of moths to detect the echolocation calls of predatory bats. Although the neurophysiology of bat detection has been intensively studied in moths for decades, the relationship between sound-induced movement of the noctuid tympanic membrane and action potentials in the auditory sensory cells (A1 and A2) has received little attention. Using laser Doppler vibrometry, we measured the velocity and displacement of the tympanum in response to pure tone pulses for moths that were intact or prepared for neural recording. When recording from the auditory nerve, the displacement of the tympanum at the neural threshold remained constant across frequencies, whereas velocity varied with frequency. This suggests that the key biophysical parameter for triggering action potentials in the sensory cells of noctuid moths is tympanum displacement, not velocity. The validity of studies on the neurophysiology of moth hearing rests on the assumption that the dissection and recording procedures do not affect the biomechanics of the ear. There were no consistent differences in tympanal velocity or displacement when moths were intact or prepared for neural recordings for sound levels close to neural threshold, indicating that this and other neurophysiological studies provide good estimates of what intact moths hear at threshold.

10aauditory threshold10aLepidoptera10amoth auditory biomechanics10aneurophysiology1 aHofstede, Hannah, M. ter1 aGoerlitz, Holger, R.1 aMontealegre-Zapata, Fernando1 aRobert, Daniel uhttp://link.springer.com/10.1007/s00114-011-0851-702036nas a2200157 4500008004100000245008500041210006900126260001600195300001200211490000600223520143000229100002401659700002401683700003301707856013801740 2017 eng d00aNon-invasive biophysical measurement of travelling waves in the insect inner ear0 aNoninvasive biophysical measurement of travelling waves in the i cJul-18-2017 a170-1710 v43 a

Frequency analysis in the mammalian cochlea depends on the propagation of frequency information in the form of a travelling wave (TW) across tonotopically arranged auditory sensilla. TWs have been directly observed in the basilar papilla of birds and the ears of bush-crickets (Insecta: Orthoptera) and have also been indirectly inferred in the hearing organs of some reptiles and frogs. Existing experimental approaches to measure TW function in tetrapods and bush-crickets are inherently invasive, compromising the fine-scale mechanics of each system. Located in the forelegs, the bush-cricket ear exhibits outer, middle and inner components; the inner ear containing tonotopically arranged auditory sensilla within a fluid-filled cavity, and externally protected by the leg cuticle. Here, we report bush-crickets with transparent ear cuticles as potential model species for direct, non-invasive measuring of TWs and tonotopy. Using laser Doppler vibrometry and spectroscopy, we show that increased transmittance of light through the ear cuticle allows for effective non-invasive measurements of TWs and frequency mapping. More transparent cuticles allow several properties of TWs to be precisely recovered and measured in vivo from intact specimens. Our approach provides an innovative, non-invasive alternative to measure the natural motion of the sensilla-bearing surface embedded in the intact inner ear fluid.

1 aSarria-S, Fabio, A.1 aSoulsbury, Carl, D.1 aMontealegre-Zapata, Fernando uhttp://rsos.royalsocietypublishing.org/lookup/doi/10.1098/rsos.170171https://syndication.highwire.org/content/doi/10.1098/rsos.17017102284nas a2200265 4500008004100000022001300041245015400054210006900208260001600277300001200293490000800305520141800313653001701731653001701748653001701765653002901782653001401811653001601825653001501841100002401856700001901880700002001899700003301919856006601952 2016 eng d a0044523100aWing mechanics, vibrational and acoustic communication in a new bush-cricket species of the genus Copiphora (Orthoptera: Tettigoniidae) from Colombia0 aWing mechanics vibrational and acoustic communication in a new b cJan-07-2016 a55 - 650 v2633 a

Male bush-crickets produce acoustic signals by wing stridulation to call females. Several species also alternate vibratory signals with acoustic calls for intraspecific communication, a way to reduce risk of detection by eavesdropping predators. Both modes of communication have been documented mostly in neotropical species, for example in the genus Copiphora. In this article, we studied vibratory and acoustic signals and the biophysics of wing resonance in Copiphora vigorosa, a new species from the rainforest of Colombia. Different from other Copiphora species in which the acoustic signals have been properly documented as pure tones, C. vigorosa males produce a complex modulated broadband call peaking at ca. 30 kHz. Since males of this species do rarely sing, we also report that substratum vibrations have been adopted in this species as a persistent communication channel. Wing resonances and substratum vibrations were measured using a μ-scanning Laser Doppler Vibrometry. We found that the stridulatory areas of both wings exhibit a relatively broad-frequency response and the combined vibration outputs fits with the calling song spectrum breadth. The broadband calling song spectrum results from several wing resonances activated simultaneously during stridulation. Under laboratory conditions the calling song duty cycle is very low and males spend more time tremulating than singing.

10abioacoustics10abiotremology10abush-cricket10alaser Doppler vibrometer10aresonance10atremulation10aultrasound1 aSarria-S, Fabio, A.1 aBuxton, Kallum1 aJonsson, Thorin1 aMontealegre-Zapata, Fernando uhttp://linkinghub.elsevier.com/retrieve/pii/S004452311630058400644nas a2200181 4500008004100000022001400041245006300055210006300118260002000181300001400201490000800215100003300223700001600256700002500272700001600297700001900313856013000332 2012 eng d a0036-807500aConvergent Evolution Between Insect and Mammalian Audition0 aConvergent Evolution Between Insect and Mammalian Audition cApr-15-20122141 a968 - 9710 v3381 aMontealegre-Zapata, Fernando1 aJonsson, T.1 aRobson-Brown, K., A.1 aPostles, M.1 aRobert, Daniel uhttp://www.sciencemag.org/cgi/doi/10.1126/science.1225271https://syndication.highwire.org/content/doi/10.1126/science.122527102623nas a2200193 4500008004100000022001400041245014900055210006900204260002000273300001600293490000800309520186800317100002202185700002402207700002002231700002202251700003302273856012302306 2017 eng d a0022-094900aFunctional morphology of tegmina-based stridulation in the relict species Cyphoderris monstrosa (Orthoptera: Ensifera: Prophalangopsidae)0 aFunctional morphology of tegminabased stridulation in the relict cDec-28-20172143 a1112 - 11210 v2203 a

Male grigs, bush crickets and crickets produce mating calls by tegminal stridulation: the scraping together of modified forewings functioning as sound generators. Bush crickets (Tettigoniidae) and crickets (Gryllinae) diverged some 240 million years ago, with each lineage developing unique characteristics in wing morphology and the associated mechanics of stridulation. The grigs (Prophalangopsidae), a relict lineage more closely related to bush crickets than to crickets, are believed to retain plesiomorphic features of wing morphology. The wing cells widely involved in sound production, such as the harp and mirror, are comparatively small, poorly delimited and/or partially filled with cross-veins. Such morphology is similarly observed in the earliest stridulating ensiferans, for which stridulatory mechanics remains poorly understood. The grigs, therefore, are of major importance to investigate the early evolutionary stages of tegminal stridulation, a critical innovation in the evolution of the Orthoptera. The aim of this study is to appreciate the degree of specialization on grig forewings, through identification of sound radiating areas and their properties. For well-grounded comparisons, homologies in wing venation (and associated areas) of grigs and bush crickets are re-evaluated. Then, using direct evidence, this study confirms the mirror cell, in association with two other areas (termed ‘neck’ and ‘pre-mirror’), as the acoustic resonator in the grig Cyphoderris monstrosa. Despite the use of largely symmetrical resonators, as found in field crickets, analogous features of stridulatory mechanics are observed between C. monstrosa and bush crickets. Both morphology and function in grigs represents transitional stages between unspecialized forewings and derived conditions observed in modern species.

1 aBĂ©thoux, Olivier1 aSarria-S, Fabio, A.1 aJonsson, Thorin1 aMason, Andrew, C.1 aMontealegre-Zapata, Fernando uhttp://jeb.biologists.org/lookup/doi/10.1242/jeb.153106https://syndication.highwire.org/content/doi/10.1242/jeb.15310602023nas a2200193 4500008004100000245016200041210006900203260001600272300001100288490000600299520131800305100002401623700002201647700002601669700002601695700003301721700002301754856005201777 2014 eng d00aShrinking Wings for Ultrasonic Pitch Production: Hyperintense Ultra-Short-Wavelength Calls in a New Genus of Neotropical Katydids (Orthoptera: Tettigoniidae)0 aShrinking Wings for Ultrasonic Pitch Production Hyperintense Ult cMay-06-2014 ae987080 v93 a

This article reports the discovery of a new genus and three species of predaceous katydid (Insecta: Orthoptera) from Colombia and Ecuador in which males produce the highest frequency ultrasonic calling songs so far recorded from an arthropod. Male katydids sing by rubbing their wings together to attract distant females. Their song frequencies usually range from audio (5 kHz) to low ultrasonic (30 kHz). However, males of Supersonus spp. call females at 115 kHz, 125 kHz, and 150 kHz. Exceeding the human hearing range (50 Hz–20 kHz) by an order of magnitude, these insects also emit their ultrasound at unusually elevated sound pressure levels (SPL). In all three species these calls exceed 110 dB SPL rms re 20 m Pa (at 15 cm). Males of Supersonus spp. have unusually reduced forewings (,0.5 mm 2 ). Only the right wing radiates appreciable sound, the left bears the file and does not show a particular resonance. In contrast to most katydids, males of Supersonus spp. position and move their wings during sound production so that the concave aspect of the right wing, underlain by the insect dorsum, forms a contained cavity with sharp resonance. The observed high SPL at extreme carrier frequencies can be explained by wing anatomy, a resonant cavity with a membrane, and cuticle deformation.

1 aSarria-S, Fabio, A.1 aMorris, Glenn, K.1 aWindmill, James, F.C.1 aJackson, Joseph, Curt1 aMontealegre-Zapata, Fernando1 aEtges, William, J. uhttp://dx.plos.org/10.1371/journal.pone.009870802155nas a2200241 4500008004100000022001400041245012900055210006900184260001600253300001200269490000700281520136500288653001701653653001301670653001901683653001701702653001501719100002601734700002001760700003201780700003301812856006801845 2014 eng d a0952-462200aUltrasonic reverse stridulation in the spider-like katydid Arachnoscelis (Orthoptera: Listrosceledinae)0 aUltrasonic reverse stridulation in the spiderlike katydid iArach cFeb-01-2014 a67 - 770 v233 a

This paper illustrates the biomechanics of sound production in the neotropical predaceous katydid Arachnoscelis arachnoides (Insecta: Orthoptera: Tettigoniidae). Described and previously known from only one male specimen, this genus of predaceous katydids resembles spiders in their general body appearance. To call distant females, male katydids produce songs by stridulation where one forewing possessing a sclerotized file rubs against a row of teeth (scraper) on the other wing. In most katydid species, the songs are produced during the wing-closing phase of the stridulation. Morphological comparative studies of the stridulatory apparatus of the type specimen of Arachnoscelis arachnoides and males of other closely related species suggest that this insect sings with a frequency of ca. 80 kHz to attract conspecific females. We found an abundant population of A. arachnoides in Central Northeast of Colombia and undertook a complete analysis of the biomechanics of stridulation in this species. Using ultrasound-sensitive equipment and high-speed video, we determined that male A. arachnoides sing at ca. 74 kHz and use elastic energy and wing deformation to reach such ultrasonic frequencies. In contrast to most katydids, males of this species produce their calls during the opening phase of the wing; this form of stridulation is discussed.

10abiomechanics10aColombia10aelastic energy10astridulation10aultrasound1 aChivers, Benedict, D.1 aJonsson, Thorin1 aCadena-Castaneda, Oscar, J.1 aMontealegre-Zapata, Fernando uhttp://www.tandfonline.com/doi/abs/10.1080/09524622.2013.81663901211nas a2200241 4500008004100000022001400041245016400055210006900219260001600288300001400304490000700318520039100325653001400716653001300730653001200743653001200755653001200767653001700779100003300796700002400829700002200853856009400875 2003 eng d a1082-646700aPanoploscelis specularis (Orthoptera: Tettigoniidae: Pseudophyllinae): extraordinary female sound generator, male description, male protest and calling signals0 aPanoploscelis specularis Orthoptera Tettigoniidae Pseudophyllina cJan-12-2003 a173 - 1810 v123 a

Females of Panoploscelis specularis present a dramatic modification of their forewings for stridulation. The female generator is illustrated and its distinct form contrasted with that of males. The physical form of the signals that females might produce is inferred; male calling and protest signals are characterized. The male of P. specularis is described for the first time.

10aacoustics10aColombia10adefense10aEcuador10akatydid10astridulation1 aMontealegre-Zapata, Fernando1 aGuerra, Patrick, A.1 aMorris, Glenn, K. uhttp://www.bioone.org/doi/abs/10.1665/1082-6467%282003%29012%5B0173%3APSOTPE%5D2.0.CO%3B201817nas a2200265 4500008004100000022001400041245012000055210006900175260001600244300001600260490000800276520100400284653002501288653001701313653001601330653001901346653001801365100001401383700003301397700001901430700001801449700001601467700001201483856005601495 2012 eng d a0027-842400aWing stridulation in a Jurassic katydid (Insecta, Orthoptera) produced low-pitched musical calls to attract females0 aWing stridulation in a Jurassic katydid Insecta Orthoptera produ cJun-03-2012 a3868 - 38730 v1093 a

Behaviors are challenging to reconstruct for extinct species, particularly the nature and origins of acoustic communication. Here we unravel the song of Archaboilus musicus Gu, Engel and Ren sp. nov., a 165 million year old stridulating katydid. From the exceptionally preserved morphology of its stridulatory apparatus in the forewings and phylogenetic comparison with extant species, we reveal that A. musicus radiated pure-tone (musical) songs using a resonant mechanism tuned at a frequency of 6.4 kHz. Contrary to previous scenarios, musical songs were an early innovation, preceding the broad-bandwidth songs of extant katydids. Providing an accurate insight into paleoacoustic ecology, the low-frequency musical song of A. musicus was well-adapted to communication in the lightly cluttered environment of the mid-Jurassic forest produced by coniferous trees and giant ferns, suggesting that reptilian, amphibian, and mammalian insectivores could have also heard A. musicus' song.

10abiological asymmetry10abiomechanics10abushcricket10acall evolution10aTettigoniidae1 aGu, J.-J.1 aMontealegre-Zapata, Fernando1 aRobert, Daniel1 aEngel, M., S.1 aQiao, G.-X.1 aRen, D. uhttp://www.pnas.org/cgi/doi/10.1073/pnas.1118372109