Thesis (PhD)--Stellenbosch University, 2014. / ENGLISH ABSTRACT: Soft, serene insect songs add an intrinsic aesthetic value to the landscape. Yet these songs also have an important biological relevance. Acoustic signals across the landscape carry a multitude of localized information allowing organisms to communicate invisibly within their environment. Ensifera are cryptic participants of nocturnal soundscapes, contributing to ambient acoustics through their diverse range of proclamation songs. Although not without inherent risks and constraints, the single most important function of signalling is sexual advertising and pair formation. In order for acoustic communication to be effective, signals must maintain their encoded information so as to lead to positive phonotaxis in the receiver towards the emitter. In any given environment, communication is constrained by various local abiotic and biotic factors, resulting in Ensifera utilizing acoustic niches, shifting species songs spectrally, spatially and temporally for their optimal propagation in the environment. Besides the importance of Ensifera songs from an ethological point of view, the multitude of species-specific signals provide an acoustic tapestry representing species diversity across ecological gradients and over time. Acoustic inventorying and monitoring of the landscape can reflect the environmental status of ecological systems, from natural to disturbed by human influence. In contrast to traditional survey techniques, sound recording and interpretation is a non-invasive method that allows for the detection and classification of highly cryptic, yet insightful indicators of ecosystem change. Here, acoustic monitoring was used across diverse ecological gradients to improve understanding of species diversity patterns, and how they change in response to both natural gradients and in response to the human dominated landscape mosaic. This study was undertaken in three geographic locations from tropical rain forest of Brunei, Borneo, to the landscapemosaic of plantation forestry in KwaZulu-Natal, South Africa, and to the botanically rich, mountain fynbos region of the Cape Floristic Region, also in South Africa. Each region provided a diverse and particular landscape to test the value of acoustic surveys for determining local diversity patterns across natural gradients and to assess the value of the technique for assessing the impact human influence across landscapes. In tropical rainforests, an entire acoustic guild was investigated to determine how acoustic species partition their acoustic communication channels spectrally, temporally and spatially, to avoid acoustic interference. The overall assemblage showed considerable spectral partitioning. Diurnally active species showed low temporal niche overlap, whereas nocturnal species did not utilize temporal partitioning. Lack of nocturnal temporal partitioning suggests other mechanisms of acoustic avoidance are sufficient to avoid acoustic overlap, or that there are insufficient cues to partition nocturnal acoustic environments. Acoustic species also utilized spatial partitioning, with distinct acoustic assemblages at vertical heights and with elevation. Utilization of a range of different strategies allow many species to communicate with conspecifics with little or no interference from other species in a signal rich environment. Acoustic profiling was also undertaken in KwaZulu-Natal, South Africa, across a plantation forestry landscape mosaic with diverse ecological gradients containing both alien and indigenous vegetation, as well as boarding large natural protected areas. Areas covered in alien timber or non-endemic grass were devoid of acoustic signals. Managed areas that were mowed and heavily grazed were not effective in maintaining the natural complement of nocturnal acoustic species. Within natural vegetation patches inside plantations, acoustic species richness increased with plant heterogeneity and patch size. Patches of indigenous vegetation within the plantation matrix effectively reduced the contrast of transformed landscapes with surrounding natural areas, with indigenous forest patches containing a highly characteristic acoustic species assemblage. Within the botanically rich, mountain fynbos region of the Cape Floristic Region, acoustic profiling was conducted across gradients of elevation, season and vegetation. Across these gradients, katydid acoustic signals were identified and characterized for the first time. This resulted in the discovery of two new katydid species and a novel sound producing structure in a carabid beetle, a species previously unknown to produce sound. Acoustic diversity across seasonal and elevational gradients increased with increasing temperatures. Climatic variability along the elevational gradient produced variation in seasonal phenology. Katydids also utilized high frequency acoustic signals, which is probably an adaptation to overcome background noise from wind, so prevalent in this area. Furthermore, despite producing conspicuous signals for mate attraction and pair formation, katydids were found not to be part of bat-eared fox diet, an insectivorous, nocturnal predator that uses its characteristic large ears to detect sounds made by invertebrate prey. This study shows the value of using acoustic emissions from katydids to identify acoustic diversity patterns across ecological gradients and in response to human impacts on the landscape. / AFRIKAANSE OPSOMMING: Sagte, rustige insekliedjies voeg ‘n intrinsieke estetiese waarde aan die landskap. Tog is hierdie liedjies ook van belangrike biologiese waarde. Akoestiese seine oor die landskap dra ‘n magdom plaaslike inligting wat organismes in staat stel om onsigbaar te kommunikeer binne hul omgewing. Langhoringsprinkane is kriptiese deelnemers van die nagtelike klankomgewing en dra by tot die omringende akoestiek deur hul verskeidenheid van proklamasieliedjies. Alhoewel dit nie sonder inherente risiko’s en beperkings is nie, is die belangrikste funksie van seine seksuele advertering en paarvorming. Vir akoestiese kommunikasie om effektief te wees, moet seine hul geënkodeerde inligting handhaaf, sodat dit sal lei tot positiewe fonotaksis in die ontvanger teenoor die emittor. In enige gegewe omgewing, word kommunikasie beperk deur verskeie plaaslike abiotiese en biotiese faktore. Dit lei tot die gebruik van akoestiese nisse deur langhoringsprinkane, wat hulle liedjies spektraal, ruimtelik en temporeel aanpas vir optimale verspreiding in die omgewing. Benewens die belang van die langhoringsprinkaan liedjies uit ʼn etologiese oogpunt, bied die menigte spesie-spesifieke seine ʼn akoestiese tapisserie wat spesiesdiversiteit verteenwoordig oor ekologiese gradiënte en oor tyd. Akoestiese opname en monitering van die landskap kan die omgewingstoestand van ekologiese stelsels weerspieël, van natuurlike tot menslik versteurde stelsels. In teenstelling met tradisionele opnametegnieke, is klankopname en interpretasie ʼn nie-indringende metode wat dit moontlik maak om hoogs kriptiese, nog insiggewende indikators van ekosisteemverandering op te spoor en te klassifiseer. In hierdie studie is akoestiese monitering gebruik oor diverse ekologiese gradiënte om ons begrip te verbeter van spesies diversiteitspatrone, en hoe dit verander in reaksie op beide natuurlike gradiënte en in reaksie op die menslik gedomineerde landskapmosaïek. Hierdie studie is onderneem in drie geografiese liggings: tropiese reënwoud in Brunei, Borneo, die landskapmosaïek van plantasiebosbou in KwaZulu-Natal, Suid-Afrika, en die plantryke, bergfynbos-streek van die Kaap Floristiese Streek, ook in Suid-Afrika. Elke streek het 'n diverse en besondere landskap verskaf om die waarde van akoestiese opnames te toets vir die bepaling van plaaslike diversiteitspatrone in natuurlike gradiënte, asook om die waarde van die tegniek te bepaal vir die beoordeling van die impak van menslike invloed oor landskappe. In tropiese reënwoude, is 'n hele akoestiese gilde ondersoek om te bepaal hoe akoestiese spesies hul akoestiese kommunikasiekanale spektraal, ruimtelik en temporeel verdeel om akoestiese inmenging te vermy. Die algehele groep het aansienlike spektrale verdeling getoon. Dagaktiewe spesies het lae temporele nisoorvleueling getoon, terwyl nagtelike spesie nie temporele verdeling benut het nie. Gebrek aan nagtelike temporele verdeling dui daarop dat ander meganismes van akoestiese vermyding voldoende is om akoestiese oorvleueling te vermy, of dat daar onvoldoende seine is om nagtelike akoestiese omgewings te verdeel. Akoestiese spesies het ook ruimtelike verdeling benut, met verskillende akoestiese spesiesversamelings op vertikale hoogtes en met hoogte bo seevlak. Die gebruik van 'n verskeidenheid van strategieë maak dit moontlik vir spesies om te kommunikeer met min of geen inmenging van ander spesies in 'n seinryke omgewing. Akoestiese profielsamestelling is ook onderneem in KwaZulu-Natal, Suid-Afrika, oor 'n plantasiebosbou landskapmosaïek met diverse ekologiese gradiënte wat beide uitheemse en inheemse plantegroei, sowel as groot, natuurlike, beskermde gebiede ingesluit het. Gebiede wat bestaan het uit uitheemse timmerhoutbome of nie-endemiese gras, was heeltemal sonder akoestiese seine. Bestuursgebiede wat gesny en swaar bewei was, het nie doeltreffend die natuurlike komplement van nagtelike akoestiese spesies gehandhaaf nie. In natuurlike plantegroei fragmente binne plantasies, het akoestiese spesiesrykheid toegeneem met plantverskeidenheid en fragmentgrootte. Fragmente van inheemse plantegroei binne die plantasiematriks het effektief die kontras van getransformeerde landskappe met omliggende natuurlike gebiede verminder, en inheemse woudefragmente het hoogs kenmerkende akoestiese spesiesversamelings gehad. Binne die plantryke bergfynbosstreek van die Kaap Floristiese Streek, was akoestiese profielsamestelling gedoen oor gradiënte van hoogte bo seevlak, seisoen en plantegroei. Oor hierdie gradiënte, is langhoringsprinkaan akoestiese seine geïdentifiseer en gekenmerk vir die eerste keer. Dit het gelei tot die ontdekking van twee nuwe langhoringsprinkaan spesies en 'n nuwe klankvervaardiging struktuur in 'n Carabid kewer, 'n spesie wat nie voorheen bekend was om klank te produseer nie. Akoestiese diversiteit oor gradiënte van seisoen en hoogte bo seevlak het toegeneem met toenemende temperature. Variasie in klimaatstoestande oor die gradiënt van hoogte bo seevlak het variasie in seisoenale fenologie veroorsaak. Langhoringsprinkane het ook hoë frekwensie akoestiese seine benut, wat waarskynlik 'n aanpassing is om agtergrondgeraas van die wind, wat so algemeen is in hierdie gebied, te bowe te kom. Verder, ten spyte van die vervaardiging van opvallende seine vir maataantrekking en paarvorming, het langhoringsprinkane nie deel gevorm van die bakoorjakkals se dieet nie. Hierdie is 'n insekvretende, nagtelike roofdier wat gebruik maak van sy kenmerkende groot ore om klanke op te spoor wat gemaak word deur invertebraatprooi. Hierdie studie toon die waarde van die gebruik van akoestiese seine van langhoringsprinkane om akoestiese diversiteitspatrone te identifiseer oor ekologiese gradiënte en in reaksie op menslike impakte op die landskap.

O objetivo deste trabalho foi elucidar aspectos morfológicos, ecológicos e comportamentais dos percevejos castanhos. Foram realizados estudos de levantamento populacional, dimorfismo sexual, polimorfismo alar e comunicação sonora com diferentes espécies de Scaptocoris. O levantamento populacional e a distribuição vertical no solo foi realizado com S. carvalhoi. As ninfas foram predominantes durante todo o período, não sendo influenciadas pela precipitação média mensal. O número de adultos foi dependente das chuvas, ocorrendo uma maior incidência destes com o incremento da precipitação. Ninfas e adultos mantiveram-se nas camadas mais superficiais do solo no período chuvoso e aprofundaram-se nos períodos de deficiência hídrica. Os adultos foram menos tolerantes ao estresse hídrico e aprofundaram-se logo no início da época seca, enquanto as ninfas o fizeram somente no final desse período. No estudo de dimorfismo sexual, as genitálias externas de machos e fêmeas de S. carvalhoi, S. buckupi e S. castanea foram extraídas e analisadas. As fêmeas se diferenciaram dos machos por apresentarem placas genitais. Nas fêmeas de S. carvalhoi o laterotergito 8 é dividido em duas partes, ao invés da placa única que forma a genitália de S. castanea e S. buckupi. Nos machos, a genitália é formada pelo 9º segmento abdominal da qual se projetam o 10º segmento abdominal e os parâmeros, sendo esta característica semelhante nas três espécies. No estudo de polimorfismo alar foram mensurados o comprimento do corpo, escutelo, asas anteriores e posteriores, comprimento da membrana, e distância entre a inserção da asa e o ápice do abdome em S. carvalhoi. Uma análise de agrupamento demonstrou a existência de dimorfismo alar, serparando-os em indivíduos de asas curtas (braquípteros) e longas (macrópteros). Nos braquípteros as asas não recobriram os últimos tergitos e nos macrópteros elas ultrapassaram o ápice do abdome. As razões entre o comprimento do corpo e das asas anteriores (RAC) e posteriores (RPC), foram superiores a 0,8 e 0,6, respectivamente, para os indivíduos macrópteros. Os braquípteros predominaram no solo durante todo o ano, apresentando menor mobilidade e asas não são funcionais. Os macrópteros ocorreram, principalmente, no início das chuvas e após um longo período seco, apresentando maior mobilidade e reação de vôo em 67% dos casos. A maior freqüência de macrópteros na camada mais superficial do solo (até 20 cm), coincidiu com a época de revoada nestes insetos e os adultos coletados após a revoada foram todos macrópteros, demonstrando a importância destes indivíduos na dispersão para novas áreas. O estudo da morfologia do aparato estridulatório localizado nos abdomes e asas posteriores de S. carvalhoi e S. castanea apresentou diferenças entre os sexos e espécies. O registro dos sons produzidos por estas estruturas e a sua dispersão através do substrato (solo e planta), confirmou que machos e fêmeas de cada espécie produzem sons distintos, e que a propagação deste som no solo ocorre somente a curtas distâncias. / The objective of this study was to elucidate morphological, ecological and behavioral aspects of the burrowing bugs (Scaptocoris). Population dynamic studies, sexual dimorphism, wing polymorphism, and acoustic communication were investigated on different species of Scaptocoris. Studies on population dynamics and vertical distribution in the soil were carried out with S. carvalhoi. Nymphs occurred during the entire length of the study, and their abundance was not influenced by monthly rainfall. Adult population augmented with the increase of rainfall. Nymphs and adults were located in the top layer of the soil during the rainy season and they were found deeper in the soil during droughts. Adults were less tolerant to droughts than nymphs. External male and female genitalia of S. carvalhoi, S. buckupi and S. castanea were extracted for sexual dimorphism studies. Females differed from males by presenting genital plates. Females of S. carvalhoi have the laterotergite 8 separated in two parts, differently from S. castanea e S. buckupi that have only one plate. Male genitalia present the 10th abdominal segment and paramers linked to the 9th segment. This characteristic is similar among the three species. The wing polymorphism was compared by measuring body length, scutellum, anterior and posterior wings, membrane length and distance from the point of insertion of the wing to the apical part of the abdomen of S. carvalhoi. Cluster analysis allowed the identification of two groups based on the wing dimorphism. It was possible to classify each group, according to wing length as brachypterous (short wings) and macropterous (long wings). Wings of brachypterous specimens do not cover the last tergites and the wings of macropterous surpass the apical part of the abdomen. The ratio of the body length and anterior wings (RAC) e posterior wings (RPC) was greater than 0.8 and 0.6, respectively, for macropterous specimens. Brachypterous insects were more prevalent on the soil than macropterous. They were less active and their wings were not functional. Macropterous stinkbugs occurred predominantly during the onset of the rainfall and after long droughts. They were more active flying in 67% of the times when touched. The higher frequency of macropterous stinkbugs in the top soil layer (first 20 cm), occurred during swarm when they were the only group of adult stinkbugs collected, demonstrating their importance on dispersion to new areas. Morphological differences of the stridulatory apparatus, located at the abdomen near the posterior wings, were detected for S. carvalhoi and S. castanea and also between males and females. The sounds produced by these insects and its propagation across the substrate (soil and plant) varied between species and sexes. The sounds propagate only within short distances.

Submitted by PPG Zoologia (zoologia-pg@pucrs.br) on 2018-09-13T17:02:53Z No. of bitstreams: 1 Disserta??o - Escalona - Mois?s.pdf: 3628906 bytes, checksum: a6fc5f6bc6015a1a1a95f67e3c5c5f16 (MD5) / Approved for entry into archive by Sheila Dias (sheila.dias@pucrs.br) on 2018-09-14T17:19:08Z (GMT) No. of bitstreams: 1 Disserta??o - Escalona - Mois?s.pdf: 3628906 bytes, checksum: a6fc5f6bc6015a1a1a95f67e3c5c5f16 (MD5) / Made available in DSpace on 2018-09-14T17:46:57Z (GMT). No. of bitstreams: 1 Disserta??o - Escalona - Mois?s.pdf: 3628906 bytes, checksum: a6fc5f6bc6015a1a1a95f67e3c5c5f16 (MD5) Previous issue date: 2018-03-28 / Conselho Nacional de Pesquisa e Desenvolvimento Cient?fico e Tecnol?gico - CNPq / Os anuros emitem cantos de an?ncio com o objetivo de atrair parceiros e repelir concorrentes co-espec?ficos. Eu investiguei a associa??o entre o comprimento do corpo, o s?tio de vocaliza??o, o cuidado parental e a varia??o em propriedades ac?sticas do canto (dura??o do canto, n?mero de notas, freq??ncia de pico, largura de banda de freq??ncia e estrutura do canto) em pererecasde-vidro?uma fam?lia de pererecas que habitam florestas neotropicais?controlando para efeitos de ancestralidade compartilhada. Eu tamb?m explorei o tempo e o modo de evolu??o dessas caracter?sticas e as comparei com as de tr?s caracter?sticas morfol?gicas (comprimento do corpo, largura da cabe?a e comprimento da t?bia). Eu gerei e compilei dados ac?sticos para 72 esp?cies de pererecas-de-vidro (46 % da riqueza descrita), incluindo representantes de todos os g?neros, e analisei os dados usando m?todos comparativos filogen?ticos. Descobri que quase todas as caracter?sticas ac?sticas avaliadas tinham sinal filogen?tico significativo, mas geralmente modesto. O pico de frequ?ncia dos cantos ? significativamente associado ao comprimento do corpo, enquanto a estrutura dos cantos ? significativamente associada ao s?tio de vocaliza??o e ao sexo do adulto que desempenha cuidado-parental. Meus resultados sustentam que, al?m das restri??es filogen?ticas, o tamanho do corpo est? associado ? evolu??o dos cantos de an?ncio em Centrolenidae. A diversifica??o de caracter?sticas ac?sticas ao longo da filogenia de pererecas-de-vidro concorda com a radia??o dos taxa na transi??o entre Mioceno e Plioceno. A disparidade estimada para caracter?sticas ac?sticas foi geralmente maior que a de caracter?sticas morfol?gicas, e o pico de disparidade de caracter?sticas ac?sticas geralmente ocorreu mais tarde na evolu??o do grupo, sugerindo um in?cio historicamente recente da pronunciada diverg?ncia ac?stica entre esp?cies deste clado. / Anurans emit advertisement calls with the purpose of attracting mates and repelling conspecific competitors. I investigated the association between body size, calling site and parental care with call variables (call duration, number of notes, peak frequency, frequency bandwidth and call structure) in glassfrogs?a family of leaf-dwelling frogs which inhabit Neotropical forests?while controlling for effects of shared ancestry. I also explored the tempo and mode of evolution of these traits and compared them with those of three morphological traits (snout-vent length, and relative head width and tibia length). I generated and compiled acoustic data for 72 glassfrog species (46 % of described diversity), including representatives of all genera (n = 12), and analyzed the data using phylogenetic comparative methods. I found that almost all acoustic traits evaluated had significant, but generally modest phylogenetic signal. Peak frequency of calls is significantly associated with body-size, while call structure is significantly associated with calling site and with the sex of the caring parent. My results support that, besides phylogenetic constraints, body size is associated with the evolution of advertisement calls in glassfrogs. Diversification of acoustic traits along the phylogeny of glassfrogs is coincident with the radiation of extant taxa at the Miocene-Pliocene boundary. The estimated disparity of acoustic traits was larger than that of morphological traits and the peak in disparity of acoustic traits generally occurred later in the evolution of glassfrogs, suggesting a historically recent outset of the pronounced acoustic divergence in this clade.

Spectrum has been a scarce commodity in RF communication. Acoustic data communication is an alternative to RF communication where data is transmitted through sound waves. In this thesis, several different aspects of acoustic data communication are investigated. A physical test setup is built where the data communication spectrum extends up to 40 kHz. Impulse response of the acoustic channel is considered in a laboratory environment. Acoustic spectrum beyond the hearing limit between 25 kHz to 35 kHz is found as a suitable band for the developed setup. Distance and multipath components are important factors, determining the communication accuracy. The physical layer for the communication system is built by taking RF Pager system as a reference. This system is also modified to improve the performance. Dierent modulation techniques are used in order to evaluate their performances for acoustic channels. BFSK, BPSK, QPSK, GMSK, OFDM, DSSS and FHSS techniques are implemented for comparison. Total and effective bit rate are considered for the overall performance evaluation of differentt modulation techniques. Several experiments are done in laboratory environment where there are several multipath components. As the distance between the transmitter and receiver is increased, path loss and multipath increases. It is shown that certain modulation techniques are more robust to multipath and are better candidates for acoustic communication. While acoustic environment is inefficient in terms of power, it is still a good candidate for communication in short distances.

O objetivo deste trabalho foi elucidar aspectos morfológicos, ecológicos e comportamentais dos percevejos castanhos. Foram realizados estudos de levantamento populacional, dimorfismo sexual, polimorfismo alar e comunicação sonora com diferentes espécies de Scaptocoris. O levantamento populacional e a distribuição vertical no solo foi realizado com S. carvalhoi. As ninfas foram predominantes durante todo o período, não sendo influenciadas pela precipitação média mensal. O número de adultos foi dependente das chuvas, ocorrendo uma maior incidência destes com o incremento da precipitação. Ninfas e adultos mantiveram-se nas camadas mais superficiais do solo no período chuvoso e aprofundaram-se nos períodos de deficiência hídrica. Os adultos foram menos tolerantes ao estresse hídrico e aprofundaram-se logo no início da época seca, enquanto as ninfas o fizeram somente no final desse período. No estudo de dimorfismo sexual, as genitálias externas de machos e fêmeas de S. carvalhoi, S. buckupi e S. castanea foram extraídas e analisadas. As fêmeas se diferenciaram dos machos por apresentarem placas genitais. Nas fêmeas de S. carvalhoi o laterotergito 8 é dividido em duas partes, ao invés da placa única que forma a genitália de S. castanea e S. buckupi. Nos machos, a genitália é formada pelo 9º segmento abdominal da qual se projetam o 10º segmento abdominal e os parâmeros, sendo esta característica semelhante nas três espécies. No estudo de polimorfismo alar foram mensurados o comprimento do corpo, escutelo, asas anteriores e posteriores, comprimento da membrana, e distância entre a inserção da asa e o ápice do abdome em S. carvalhoi. Uma análise de agrupamento demonstrou a existência de dimorfismo alar, serparando-os em indivíduos de asas curtas (braquípteros) e longas (macrópteros). Nos braquípteros as asas não recobriram os últimos tergitos e nos macrópteros elas ultrapassaram o ápice do abdome. As razões entre o comprimento do corpo e das asas anteriores (RAC) e posteriores (RPC), foram superiores a 0,8 e 0,6, respectivamente, para os indivíduos macrópteros. Os braquípteros predominaram no solo durante todo o ano, apresentando menor mobilidade e asas não são funcionais. Os macrópteros ocorreram, principalmente, no início das chuvas e após um longo período seco, apresentando maior mobilidade e reação de vôo em 67% dos casos. A maior freqüência de macrópteros na camada mais superficial do solo (até 20 cm), coincidiu com a época de revoada nestes insetos e os adultos coletados após a revoada foram todos macrópteros, demonstrando a importância destes indivíduos na dispersão para novas áreas. O estudo da morfologia do aparato estridulatório localizado nos abdomes e asas posteriores de S. carvalhoi e S. castanea apresentou diferenças entre os sexos e espécies. O registro dos sons produzidos por estas estruturas e a sua dispersão através do substrato (solo e planta), confirmou que machos e fêmeas de cada espécie produzem sons distintos, e que a propagação deste som no solo ocorre somente a curtas distâncias. / The objective of this study was to elucidate morphological, ecological and behavioral aspects of the burrowing bugs (Scaptocoris). Population dynamic studies, sexual dimorphism, wing polymorphism, and acoustic communication were investigated on different species of Scaptocoris. Studies on population dynamics and vertical distribution in the soil were carried out with S. carvalhoi. Nymphs occurred during the entire length of the study, and their abundance was not influenced by monthly rainfall. Adult population augmented with the increase of rainfall. Nymphs and adults were located in the top layer of the soil during the rainy season and they were found deeper in the soil during droughts. Adults were less tolerant to droughts than nymphs. External male and female genitalia of S. carvalhoi, S. buckupi and S. castanea were extracted for sexual dimorphism studies. Females differed from males by presenting genital plates. Females of S. carvalhoi have the laterotergite 8 separated in two parts, differently from S. castanea e S. buckupi that have only one plate. Male genitalia present the 10th abdominal segment and paramers linked to the 9th segment. This characteristic is similar among the three species. The wing polymorphism was compared by measuring body length, scutellum, anterior and posterior wings, membrane length and distance from the point of insertion of the wing to the apical part of the abdomen of S. carvalhoi. Cluster analysis allowed the identification of two groups based on the wing dimorphism. It was possible to classify each group, according to wing length as brachypterous (short wings) and macropterous (long wings). Wings of brachypterous specimens do not cover the last tergites and the wings of macropterous surpass the apical part of the abdomen. The ratio of the body length and anterior wings (RAC) e posterior wings (RPC) was greater than 0.8 and 0.6, respectively, for macropterous specimens. Brachypterous insects were more prevalent on the soil than macropterous. They were less active and their wings were not functional. Macropterous stinkbugs occurred predominantly during the onset of the rainfall and after long droughts. They were more active flying in 67% of the times when touched. The higher frequency of macropterous stinkbugs in the top soil layer (first 20 cm), occurred during swarm when they were the only group of adult stinkbugs collected, demonstrating their importance on dispersion to new areas. Morphological differences of the stridulatory apparatus, located at the abdomen near the posterior wings, were detected for S. carvalhoi and S. castanea and also between males and females. The sounds produced by these insects and its propagation across the substrate (soil and plant) varied between species and sexes. The sounds propagate only within short distances.

No description available.

Si le signalement du prédateur provoque le plus souvent la fuite des proies, il induit parfois un comportement particulier incitant la proie à s’approcher du prédateur et le harceler pour provoquer son départ plutôt que de se mettre hors de sa portée. Ce comportement de harcèlement s’accompagne de l’émission d’un signal hétérospécifique conduisant de nombreuses espèces à venir harceler le prédateur. L’objectif de mon travail de thèse est de comprendre comment est régi le transfert d’informations entre plusieurs espèces de passereaux dans le cas du comportement de harcèlement d’un prédateur. Mes travaux montrent que le comportement de harcèlement des passereaux face à un rapace nocturne dépend du risque de prédation posé par ce prédateur. Par ailleurs, les résultats indiquent un transfert d’informations entre les espèces et mettent en évidence une propension variable des différentes espèces à se rallier autour du harceleur. Les variations observées dans la réponse aux cris de harcèlement émis par des individus hétérospécifiques peuvent dépendre de la similarité acoustique, des relations interspécifiques et des variations saisonnières. Mes résultats indiquent aussi que la connaissance préalable des signaux de harcèlement n'est pas indispensable pour induire une réponse, même si un processus d’apprentissage associatif favorise sa mise en place. Mon travail suggère également une évolution convergente des cris de harcèlement, générant des signaux dont la structure permet une localisation rapide de l’émetteur, indispensable pour rameuter des proies potentielles lors du harcèlement. L’ensemble de ces avancées nous oblige désormais à considérer la communication acoustique chez les passereaux en prenant en compte le risque de prédation, les interactions hétérospécifiques et la complexité des signaux acoustiques / Signaling the presence of a predator most often causes the escape of prey, but it sometimes induces a particular behaviour prompting prey to approach and harass the predator to cause his departure. This mobbing behaviour is associated with the emission of signals leading individuals from different species to come harass the predator. The objective of this thesis is to understand how the transfer of information between several passerine species is organized in mobbing behaviour against predators. My work shows that the mobbing behaviour of passerine birds against a nocturnal raptor depends on the predation risk imposed by this predator. In addition, my results indicate a transfer of information between species and highlight a variable propensity of different species to rally around the harasser. The observed variations in the response to heterospecific calls depended on acoustic similarity, interspecific relationships and seasonality. My results also indicate that prior knowledge of harassment signals is not essential to induce a response, even if an associative learning process promotes its implementation. My work also suggests a convergent evolution in mobbing calls, generating signals with a structure that allows the emitter to be quickly located, an essential parameter to rally potential prey during harassment. My thesis consequently shows that to better understand acoustic communication in passerine birds, it is necessary to consider predation risk, heterospecific interactions and the complexity of acoustic signals

Sound does not transmit well across the interface of two media. Therefore, most organisms communicate using one medium. Some anurans vocalize at the interface of air and water, though reception of these vocalizations is generally unknown. The túngara frog ( Engystomops pustulosus ) may be the first anuran to have evidence suggesting simultaneous acoustic communication both above and below the air-water interface. This thesis addresses whether the female túngara frog would be receptive to underwater acoustic signals and if males project their advertisement calls at biologically relevant intensities underwater. Females floated and swam with their eardrums and body walls constantly submerged. Using laser Doppler vibrometry, peak vibrations of female eardrums were found to be centered at about 3.5 kHz in air, but dropped to about 1.4 kHz underwater. The peak velocity of the eardrum was about 0.2 mm/s in air and 0.04 mm/s in water when stimulated with tones at 80 dB relative to 20 µPa. Males projected their advertisement calls with a sound pressure level of 121 dB (at 10 cm, re. 20 µPa) in water and 98 dB (at 10 cm, re. 20 µPa) in air. In relation to air, the dominant frequency of the advertisement call (0.8 kHz) was the most intense spectral band underwater whereas the dominant frequency of the chuck (2.5 kHz in air) was less intense. The advertisement signal for the male túngara frog was broadcasted underwater with more energy than in air at its main frequencies. Female eardrums were sensitive to frequencies within the male advertisement call both in air and in water, if the frequencies were transmitted at amplitudes plausible to be encountered in nature. These results strengthen the available evidence of underwater communication, and indicate the presence of auditory specializations in the acoustic communication of this species.

Background noise is ubiquitous and can impair acoustic communication and influence signaling behavior in animals. Despite evidence demonstrating myriad effects of anthropogenic noise on animal communication, precisely how natural background noise influences communication and behavior remains unclear. Yet, natural sources of background noise, such as rushing rivers or crashing ocean surf, share similar power spectra to sources of anthropogenic noise and can occur at high amplitudes, potentially masking acoustic signals. To investigate the effects of water-generated noise on songbird behavior, we experimentally broadcast landscape-level playbacks of ocean surf and river noise in coastal California, USA, and riparian habitat in Idaho, USA, respectively. In Chapter 1, we conducted a conspecific playback experiment examining how territorial defense behaviors of lazuli buntings (Passerina amoena) and spotted towhees (Pipilo maculatus) vary in response to broadcast water noise. We also incorporated cicada noise from a serendipitous Okanagana spp. emergence as a biotic source influencing lazuli bunting behavior. Both species produce songs that share substantial spectral overlap with low-frequency, water-generated noise, and lazuli bunting song shares an additional high-frequency overlap with cicada calls. Thus, there is potential for background acoustic conditions to mask conspecific signals. We found that detection and discrimination of conspecific playback occurred more slowly for both species as sound level increased. Lazuli buntings also exhibited divergent flight behaviors in response to high- and low-frequency acoustic sources, both dependent and independent of sound level. In Chapter 2, we investigated how amplitude and frequency of water-generated noise influences spectral and temporal song characteristics in six songbird species. We recorded individuals defending territories across 37 sites, with each site representing one of four acoustic environments: naturally quiet ‘controls’, naturally noisy ‘positive controls’ adjacent to the ocean or a whitewater river, ‘phantom’ playback sites with continuous broadcast of low-frequency water noise, and ‘shifted’ playback sites with continuous broadcast of high-frequency water noise. We predicted that all individuals exposed to ‘positive control’, ‘phantom’, or ‘shifted’ noise would adjust song structure, but the magnitude of signal modification would be larger in noisier locations and the type of modification would depend on the spectral profile of the acoustic environment. No two species altered songs in precisely the same way. However, song structure of all six species varied with amplitude and/or frequency of background noise. Together our results demonstrate that natural noise can impair agonistic behaviors and influence vocal structure. These findings suggest that the natural acoustic environment shapes acoustic communication, highlighting natural soundscapes as an under-appreciated axis of the environment.

L'un des plus grands défis posés par la communication est que l'information codée par l'émetteur est toujours modifiée avant d'atteindre le récepteur, et que celui-ci doit traiter cette information altérée afin de recouvrer le message. Ceci est particulièrement vrai pour la communication acoustique, où la transmission du son dans l'environnement est une source majeure de dégradation du signal, ce qui diminue l'intensité du signal relatif au bruit. La question de savoir comment les animaux transmettent l'information malgré ces conditions contraignantes a été l'objet de nombreuses études, portant soit sur l'émetteur soit sur le récepteur. Cependant, une recherche plus intégrée sur l'analyse de scènes auditives est nécessaire pour aborder cette tâche dans toute sa complexité. Le but de ma recherche était d'utiliser une approche transversale afin d'étudier comment les oiseaux s'adaptent aux contraintes de la communication à longue distance, en examinant le codage de l'information au niveau de l'émetteur, les dégradations du signal acoustiques dues à la propagation, et la discrimination de cette information dégradée par le récepteur, au niveau comportemental comme au niveau neuronal. J'ai basé mon travail sur l'idée de prendre en compte les problèmes réellement rencontrés par les animaux dans leur environnement naturel, et d'utiliser des stimuli reflétant la pertinence biologique des problèmes posés à ces animaux. J'ai choisi de me focaliser sur l'information d'identité individuelle contenue dans le cri de distance des diamants mandarins (Taeniopygia guttata) et d'examiner comment la signature vocale individuelle est codée, dégradée, puis discriminée et décodée, depuis l'émetteur jusqu'au récepteur. Cette étude montre que la signature individuelle des diamants mandarins est très résistante à la propagation, et que les paramètres acoustiques les plus individualisés varient selon la distance considérée. En testant des femelles dans les expériences de conditionnement opérant, j'ai pu montrer que celles-ci sont expertes pour discriminer entre les signature vocales dégradées de deux mâles, et qu'elles peuvent s'améliorer en s'entraînant. Enfin, j'ai montré que cette capacité de discrimination impressionnante existe aussi au niveau neuronal : nous avons montré l'existence d'une population de neurones pouvant discriminer des voix individuelles à différent degrés de dégradation, sans entrainement préalable. Ce niveau de traitement évolué, dans le cortex auditif primaire, ouvre la voie à de nouvelles recherches, à l'interface entre le traitement neuronal de l'information et le comportement / In communication systems, one of the biggest challenges is that the information encoded by the emitter is always modified before reaching the receiver, who has to process this altered information in order to recover the intended message. In acoustic communication particularly, the transmission of sound through the environment is a major source of signal degradation, caused by attenuation, absorption and reflections, all of which lead to decreases in the signal relative to the background noise. How animals deal with the need for exchanging information in spite of constraining conditions has been the subject of many studies either at the emitter or at the receiver's levels. However, a more integrated research about auditory scene analysis has seldom been used, and is needed to address the complexity of this process. The goal of my research was to use a transversal approach to study how birds adapt to the constraints of long distance communication by investigating the information coding at the emitter's level, the propagation-induced degradation of the acoustic signal, and the discrimination of this degraded information by the receiver at both the behavioral and neural levels. Taking into account the everyday issues faced by animals in their natural environment, and using stimuli and paradigms that reflected the behavioral relevance of these challenges, has been the cornerstone of my approach. Focusing on the information about individual identity in the distance calls of zebra finches Taeniopygia guttata, I investigated how the individual vocal signature is encoded, degraded, and finally discriminated, from the emitter to the receiver. This study shows that the individual signature of zebra finches is very resistant to propagation-induced degradation, and that the most individualized acoustic parameters vary depending on distance. Testing female birds in operant conditioning experiments, I showed that they are experts at discriminating between the degraded vocal signatures of two males, and that they can improve their ability substantially when they can train over increasing distances. Finally, I showed that this impressive discrimination ability also occurs at the neural level: we found a population of neurons in the avian auditory forebrain that discriminate individual voices with various degrees of propagation-induced degradation without prior familiarization or training. The finding of such a high-level auditory processing, in the primary auditory cortex, opens a new range of investigations, at the interface of neural processing and behavior