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Dissertation DISSERTATION Titel der Dissertation Multimodal signals in anurans: The role of acoustic and visual signals in the communication of foot-flagging frogs (Multimodale Signale bei Anuren: Die Rolle von akustischen und visuellen Signalen in der Kommunikation von Winkerfröschen) Verfasserin Mag. Doris Preininger angestrebter akademischer Grad Doktorin der Naturwissenschaften (Dr.rer.nat.) Wien, 5. Oktober 2012 Studienkennzahl lt. Studienblatt: A 091 439 Dissertationsgebiet lt. Studienblatt: Dr.-Studium der Naturwissenschaften, Zoologie Betreuerin / Betreuer: Ao. Univ.-Prof. Dr. Walter Hödl Acknowledgements There are many people who contributed to this thesis and I am grateful for the help, co- operation and collaboration of each one of them, but it all started with the idea and enthusi- asm of one person, who saw 15 years ago, on one of his many excursions, something very simple and extremely striking – a frog that foot flags. The frog was from the genus Staurois, the attentive observer was Walter Hödl, the latter I deeply thank for his guidance of every aspect of my work and for putting me in charge of his research idea without thinking twice. I particularly thank Markus Böckle for so many things that it would fill several pages. We started out as colleagues during our combined master studies became close friends and to this day he supports every scientific and not so scientific decision of mine. I am extremely grateful for the help, patience and friendship of Marc Sztatecsny and Iris Starnberger, they have not only enriched this project but also my life. I thank my collaborators Ulmar Grafe from Brunei and K.V. Gururaja “Guru”, S.P. Vijaya- kumar “Vijay” from India for their endless help and support and for making my field trips an exceptional experience. I thank Dagmar Schratter, Anton Weissenbacher, Regina Riegler, Evi Karell and several zoo-keepers for devoting their help and time to a new breeding and research project in the Vienna Zoo. I am especially grateful for the devotion of Thomas Wampula to this project, for his assistance in the field, for his friendship and for continuously ensuring me that everything is going to work out. I thank my parents for their encouragement when I excitedly told them: I am going to study frogs that wave their legs to communicate in tropical rainforests at the other end of the world. And although I am not sure they were as excited as I was, mainly about the “other end of the world”-part, they have supported me on every step of the way. I thank my husband for believing in me, for listening attentively for hours and hours to stories about frogs and for sometimes reminding me to turn off my computer. Since so many people have contributed to my thesis I believe it is only fair to address most sections of my introduction and conclusion as our work rather than my work! Table of content Summary ...................................................................................................................... 1 Zusammenfassung ……………………………………………………………………......... 2 Preface …………………………………………………………………………...…….…..... 3 Synopsis of the publications …...................................................................................... 9 Chapter 1. Multimodal communication in a noisy environment: A case study of the Bornean rock frog Staurois parvus ………………………………….................... 13 2. Multimodal signaling in the Small Torrent Frog (Micrixalus saxicola) in a complex acoustic environment …………………………………………………… 23 3. Divergent receiver responses to components of multimodal signals in foot-flagging frog species offer clues to visual signal evolution …………......... 47 4. The conservation breeding of two foot-flagging frog species from Borneo, Staurois parvus and Staurois guttatus ……………………………...................... 73 Concluding Discussion ………………………………………………………...……........... 85 References ………………………………………………………………………………....... 91 Appendix A. Micrixalus saxicola a foot-flagging frog from India: Acoustic and visual signaling behavior during male-male agonistic interactions……………............ 97 B. Females do have a say in the matter: Female and male vocalizations and laryngeal structures in the foot-flagging frog species Staurois guttatus ........... 103 C. Don’t get the blues: conspicuous nuptial colouration of male moor frogs (Rana arvalis) supports visual mate recognition during scramble competition in large breeding aggregations ……………...........…..................... 113 Curriculum Vitae …………………………………………………………………………...... 121 1 Summary The predominant communication mode of anuran amphibians are vocalizations, however intraspecific communication may involve multimodal (acoustic and visual) cues or signals in many more species than previously thought. Visual signals may act as an additional or a complementary mode of communication in noisy environments. Foot flagging as a striking form of visual signaling behavior has evolved in at least in 16 species from 5 different fami- lies mainly living along fast-flowing streams, which generate continuous broadband back- ground noise. To better understand the function of foot flagging as multimodal signal compo- nent, we studied three Asian species from two families and performed cue-isolation experi- ments in the field. The Bornean species Staurois parvus and S. guttatus avoid acoustic interference of am- bient stream noise by using call frequencies less masked by the background and utilize ac- companying visual signals to announce the readiness to defend calling sites, interestingly this behavior can be observed already in juveniles bred in the Vienna Zoo. Micrixalus saxicola from the Western Ghats (India) occurs along less noisy streams and acoustic signals are rather masked by chorus noise from conspecifics than by abiotic noise. Males use a variety of visual signals including foot-flagging and tapping during agonistic be- havior. Receiver responses from acoustic- vs. multimodal playback presentations provide evidence that the vocal sac acts as an additional visual cue. Comparisons of signal response to acoustic, visual and multimodal stimuli and analysis of signal brightness of foot webbings in M. saxicola and S. parvus further highlight differences in the magnitude and significance of signaling behavior in the respective species. Together these investigations allow us to draw conclusions on signal efficacy and function and help to better understand the evolution of multimodal communication in anuran. 2 Zusammenfassung Frösche kommunizieren vorwiegend mit akustischen Signalen, allerdings verwenden ei- nige Arten multimodale (akustische und visuelle) Signale während der intra-spezifischen Kommunikation. Visuelle Signale können in geräuschvoller Umgebung eine zusätzliche oder ergänzende Kommunikationsweise ermöglichen. Das Beinwinken (foot flagging) eine auffäl- lige Form von visuellem Signalverhalten hat sich in mindestens 16 Arten aus 5 verschiede- nen Familien entwickelt. Die Vielzahl dieser Arten lebt und reproduziert entlang schnell- fliessender Bergbäche, die ein kontinuierliches Hintergrundrauschen generieren. Um die Funktion des Beinwinkens als multimodale Signalkomponente besser zu verstehen, haben wir drei asiatische Arten aus zwei Familien untersucht und die akustischen und visuellen Signale in Feldexperimenten getestet. Die auf Borneo heimischen Arten Staurois parvus und S. guttatus verwenden hoch- frequente Rufe und reduzieren dadurch akustische Interferenzen vom umgebenden Bach- rauschen und benutzen zusätzliche visuelle Signale, um die Verteidigungsbereitschaft eines Rufstandort anzuzeigen. Interessanter Weise kann dieses Verhalten bereits bei juvenilen Tieren, die im Wiener Tiergarten Schönbrunn gezüchtet wurden, beobachtet werden. Micrixalus saxicola aus den Western Ghats (Indien) bewohnt weniger geräuschvolle Bäche, welche die akustische Kommunikation nicht einschränken. Die akustischen Signale können jedoch von arteigenen Rufchören maskiert werden. Die Männchen verwenden visuelle Sig- nale, wie Beinwinken und Beinheben während agonistischen Interaktionen. Reaktionen auf akustische oder multimodale Signalstimuli zeigen, dass die Schallblase als zusätzliches vi- suelles Hinweissignal fungiert. Gegenüberstellungen von Reaktionen auf akustische, visuelle und multimodale Stimuli und Analysen der Signalhelligkeit der Schwimmhäute von M. saxico- la und S. parvus, verdeutlichen die Unterschiede in Ausmaß und Stellenwert des Signalver- haltens der jeweiligen Art. Zusammengefasst ermöglichen diese Untersuchungen Schluss- folgerungen über die Funktion und Wirksamkeit der visuellen Signale in der Kommunikation von Fröschen. 3 Preface In animal communication a signal from one organism stimulates the sensory system and influences a behavioral change of a perceiver, and generally signaler and perceiver benefit from this exchange (Rendall et al. 2009; Ruxton & Schaefer 2011). On the signaler's side, signals (traits evolved for communication) are produced and correlate with an attribute of the signaler, whereas on the perceiver's side signals have to be correctly received, processed and decided upon. The production and reception of a signal is additionally influenced by the environment through which a signal travels (Endler 1992; Endler 1993b). A change in one process (production, transmission through certain environmental conditions or reception) will eventually alter the other processes and lead to evolutionary changes in signaling (Endler 2000). To understand selection pressures acting on signals and to further explain signal evolu- tion, we ask two elementary
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