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Communication and Sexual Selection in the Barking Gecko(Ptenopus

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Communication and Sexual Selection in the Barking Gecko(Ptenopus Communication and Sexual Selection in the Barking Gecko (Ptenopus kochi) Daniel Adam Polakow Town Submittedfor fulfilment Capeofthe degree Master ofScience (MSc)of University Department ofZoology University of Cape To'!Il ' •I ,, " ;r ,.:;~,~.~~- r ' • • 'l•ob ' "'' , I I' ~" • 1 • 'l " f, .t ..h or. • (' - - .... ~ , Jo - - ;:. __ ...........,. ·~· ·~~ ·~ ·~, ·--..:. :.!-r:' . ..;c: -: .. ~ . ? ...~. ~·::. The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town To Joshua Julian, Inessa Triton, Peter, Apollo Russell, Lisa, Danielle, Pan and Nicolas - the most beautiful people in my world. ' Won 't you help to sing, these songs offreedom ... ' Bob Marley Redemption Song (1980) 11 Abstract This study focused on elucidating the functional significance of some aspects of the behaviour of Koch's barking gecko, Ptenopus kochi, during a field season conducted at the Desert Ecology Research Unit in Namibia for four months in 1995. Ptenopus kochi is a terrestrial species, and males were observed calling from their burrow entrances in the' dry Kuiseb river bed during the hot sununer months. First, aspects of competition among calling males were investigated. Calling males were seen to be non-randomly distributed relative to one another with evidence for regularity of spacing in dense aggregations. Sound intensity was investigated as the mechanism of spacing, and was mathematically modelled to gauge how the intensities of the calls of nearest-neighbour males overlapped. Results revealed a constancy at which neighbouring call intensities overlap, and playback experiments ratified the presence of a critical intensity threshold in the population, beyond which male tolerance broke down. These results suggested that spacing in P. kochi is vocally-mediated and that sound intensity acts as the proximate cue through which individuals gauge the closeness of neighbouring males. Second, the inter-relationships between call features ofP . kochi's vocalisation, and how these features related to both intrinsic (morphological) and extrinsic (ambient) sources of variability were analytically explored. The principles of information theory and individual recognition by voice were used to gauge the variation manifest in the features ofP . kochi's call. A novel statistical method for disentangling the influence of incidental sources of variation from 'signal' features in animal voices was developed. Results showed how once the predictable covariance in call structure was removed, the calls ofP. kochi males were rendered statistically indistinguishable from one another. Several descriptor measures ofP . kochi's vocalisation were seen to be highly dependent on meteorological determinants, as was nightly chorus duration. Lastly, mate-choice in P. kochi was explored using correlational evidence. Results revealed that mating success was closely linked to individual male differences in display effort and chorus site burrow activity, and less to morphological or territorial characteristics. Counter-intuitively, a positive relationship between call rate and chorus site residency was found in males, suggesting a possible role for classical condition­ dependence in the evolution of female mate preference. Aspects of mate choice, together with the emergent importance of residency in this study were discussed and some ideas suggested for further research. Ul Declaration I declare that this thesis is my own work. It is beingsubmitted for thedegree of Master of Scienceat the UniversityCape of Town, SouthAfrica . It has notbeen submitted before for any degreeor examination at any other. university Signature Removed DanielAdam Polakow lV Acknowledgements Mom and Dad for your continued succour. My capricious pursuits would not be possible without your unfailing support and love. Thank you. My supervisors: Jenny Jarvis for your cardinal support and much appreciated encouragement over the last two years. Michael Cherry, too, for your support and for your helpful advice. Steven Telford, for not only being an impressive supervisor, but also for being a very special friend. Thanks Steve. The Foundation for Research and Development and the University of Cape Town for much appreciated financial support. Mary Seely and the Desert Research Foundation of Namibia for being there with all the logisitical sustenance and the Ministry of Environment and Tourism, Namibia for their assistance. To the Gobabeb endemics -Karla du Preez, Derick du Toit, Hilde Gevers, Lilo Nghinanwame, Susanne Peters, Teresa Sguazzin and Juliane Zeidler. Thanks for the company girls. Vihlo 'Snake' Mtuleni - for the most beautifully humbling perspective on the world. Thanks Snake, and good luck. Frank Echardt for the tremendous artery-clogging fry-ups every morning, afternoon and evening for several months in the desert. Thanks also for waking me at 2.30 a.m on those freezing mornings - when your enthusiasm got the better of your good judgement - to watch the fog roll in and to witness the snow. Rudiger and Ruth Rossl for the most superb introductions to the joy of German cooking, handrearing and loving fluffy toys, and the art of uncontrollable sweetness and cheerfulness. Thanks Rudy for translating bits of the Gennan literature for me. Lastly, to the BBC camera team for the laughs. So sorry we had to witness your eating of the 'gravel-plains Rossl' jam and liquorice cake. I hope that both your stomachs and senses of humour recover. v Dave Tilch for the acoustical engineering, for the anechoic chamber design, the sound blaster advice, for the linear algebra checks and for the not so linear Tsitsikarruna adventure. Ian Davidson for 'timeous' electronic skills, Shimrit Lahav-Ginott, Gary Davies and Lorraine Egan for valuable field assistance. Rupert Marshall for the Bayesian foresight. Special thanks to Wolf Haacke for kindly providing a comprehensive set of reprints on the Plenopus work, and for the useful discussion. Patricia Backwell, Michael Beecher, Rob Brooks, Neil Caithness, Mike Cherry, Mark Cooper, Derick du Toit, Jermy Jarvis, Mike Jennions, Bill Hamilton, Kate Lessells, Justin O'Riain, Paul Sherman, Andrew Spinks, and Steve Telford for some very valuable discussion and input. Bill Hamilton, Andrew Scherrer, Yehudah Werner and Gerry Wilkinson for the encouragement. Douglas Hawkins - thanks for listening so patiently from so far and so high to my multidimensional covariate-adjusted ideas, and for so perspicaciously transforming them into a statistically acceptable distribution. Graham Alexander, Michael Beecher, Sharon Doherty, Neville Eden, Mr. Jongens, Mervin Mason - for being there. Permy Beamish and Janine Dunlop for the reference hunting .. Hazel Conin, Colette Franz, Onno Huyser, Quinton Johnsone,. Harry Kentrotus, Rael Loon, Lance Myers, Throshni Naidoo, Lorenzo Prendini, Andrew Pringle, Michele Sanders, Sidney Seftel, Sandra Schakowski, Pippa Skotnes, Amy Spriggs, my UWC undergraduate class, Tracey Wallace and Anat Yakuel for making my Cape Town memories all the more unforgettable. Sheila Rock - for the colour and the smiles. Thanks Sheila! Mark 'Phredrus' Cooper. Justin O'Riain, Andrew Spinks and Laura-Anne van der Westhuizen - mostly for the tolerance, but also for the inspiration. Colin, Bertrand, Stanislaw, Alan, Daniel, Douglas, but mostly Kikis, Charles, and Rudolf for the thought. Everard, Kahlil, Will.iam and John for the poetry. Tom, Nick, Bob, Ismael, and Ludwig for the music. Club Logos for the dance. Ted and Rose for the food. Twinings for the tea. Lightspeed Inc. for the modem. Polyalphabetical ciphers for the timeless amusement. And, of course, the Mother City with your mountains, rivers, oceans and lights. V1 List of Tables Table 2.1 Results of playback experiments to males 49 Table 3.1 Literature survey of analytical aspects of vocal signatures 105 Table 3.2 Literature survey of experimental aspects of vocal signatures 111 Table 3.3 Predictor variables used in GLM analyses 116 Table 3.4 GLM result summary 117 Table 3.S Discriminant function analysis summary (naked mole-rat) 118 Table 3.6 Standardised coefficients of DF A analysis (naked mole-rat) 119 Table 3.7 Discriminant function analysis summary (barkjng gecko) 120 Table 3.8 Standardised coefficients ofDFA analysis (barking gecko) 121 Table 3.9 Chorus duration multiple-regression sununary 122 Table 5.1 Historical taxonomic features of Ptenopus 162 Vll List of Plates and figures Plate 1.1 Photograph of study site 20 Plate 1.2 Photograph of male P. kochi 21 Figure 1.1 Distribution of Ptenopus species in southern Africa 22 Figure 2.1 Mapping ofindividuals in two-dimensional space 50 Figure 2.2 Histogram of squared nearest neighbour distances 51 Figure 2.3 Observed vs. random distributions of variate y 52 Figure 2.4 Bivariate kernel plot of gecko concentration on study site 53 Figure 2.5 Bivariate kernel plot of female concentration on study site 54 Figure 2.6 Sonagram of barking gecko advertisement call 55 Figure 2.7 Sound field investigation setup 56 Figure 2.8 Observed vs. expected sound pressure decay over distance 57 Figure 2.9 Frequency distribution and coefficient of variation of r' 58 Figure 2.10 Plot of residency and fX<•> 59 Figure 2.11 Plot of fX( t) and adjusted residency scores 60 Figure 2.12 Frequency distribution of Ci Ci: 61 Figure 3.1 Sonagram of barking gecko and call descriptor measures 123 Figure 3.2 Histogram of call click number for P. kochi 124 Figure 3.3 Sonagram of naked mole-rat soft chirp and descriptors 125 Figure 3.4 Canonical plots ofDFAs (naked mole-rat) 126 Figure 3.5 Canonical plots ofDFAs (barking gecko) 127 Figure 4.1 Three-dimensional plots of residency on burrow patterning 146 Figure 4.2 Three-dimensional plots of mass on burrow patterning 147 Figure 4.3 Three-dimensional plots of SPL on burrow patterning 148 Figure 4.4 Three-dimensional plot ofmated male localities 149 Plate 5.1 Photographs of toe scaling ofP.
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