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Rhinolophus Clivosus (Chiroptera: Rhinolophidae): an Experimental Approach

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Rhinolophus Clivosus (Chiroptera: Rhinolophidae): an Experimental Approach View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Cape Town University OpenUCT The role of echolocation in communication in a high duty cycle echolocating bat, Rhinolophus clivosus (Chiroptera: Rhinolophidae): An experimental approach. By Robert Raw Thesis presented for the degree of UniversityDoctor ofof Philoso Capephy Town Department of Biological Sciences University of Cape Town September 2016 Supervised by: David Jacobs and Anna Bastian 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 I dedicate this thesis to my loving wife, Alicia Thomas-Raw i Table of Contents Declaration iii Figure Index iv Abstract viii Acknowledgements x Chapter 1 1 Chapter 2 16 2.1 Study Area 16 2.2 Study Animals 17 2.3 Animal Capture and Husbandry 22 2.4 Preparation of Playback Calls 24 2.5 Habituation – Dishabituation Playback Experiments 25 Chapter 3 33 3.1 Introduction 33 3.2 Methods for Species Discrimination Trials 36 3.3 Results for Species Discrimination Trials 40 3.4 Discussion – Species Discrimination by R. clivosus using echolocation 48 Chapter 4 53 4.1 Introduction 53 4.2 Methods for Call Component Trials 55 4.3 Results for Call Component Trials 58 4.4 Discussion – Bats use multiple components of a signal for discrimination 66 Chapter 5 72 5.1 Introduction 72 5.2 Methods for Preference Trials 77 5.3 Results for Preference Trials. 84 5.4 Discussion – Preference for gender and body condition 92 Chapter 6 101 Chapter 7 107 ii Declaration I, Robert Nicholas Vause Raw, hereby declare that the work on which this thesis is based is my original work (except where acknowledgements indicate otherwise) and that neither the whole work nor any part of it has been, is being, or is to be submitted for another degree in this or any other university. I authorise the University to reproduce for the purpose of research either the whole or any portion of the contents in any manner whatsoever. This thesis has been submitted to the TurnItIn module and I confirm that my supervisors have seen my report and any concerns revealed by such have been resolved with my supervisors. Name: Robert Raw Student Number: RWXROB003 Signature: Date: 05/09/2016 iii Figure Index Figure 2.1 Cross Section of De Hoop Guano Cave (Adapted from McDonald et al. (1990b)). Numbers 1-7 represent site of temperature measurement (Laycock, 1983). Red triangles represent roost locations for Rhinolophids observed by McDonald et al. (1990b). Blue triangle represents primary location of R. clivosus capture for this study. ................................................................................ 17 Figure 2.2 Geoffroy’s Horseshoe Bat, Rhinolophus clivosus ................................................................. 18 Figure 2.3 Map of southern Africa showing the distribution of the three horseshoe bat species used in this study: R. clivosus (Rcl, dotted line, light grey; (Monadjem et al., 2010)), R. blasii (Rbl, dashed line, dark grey; (Monadjem et al., 2010)) and R. capensis (Rca, solid line, white; (Odendaal et al., 2014)). The study site, De Hoop Nature Reserve, is indicated by a red dot. Numbers indicate the resting frequencies of the three species. ....................................................................................... 19 Figure 2.4 Blasius’s Horseshoe Bat, Rhinolophus blasii ........................................................................ 20 Figure 2.5 Cape Horseshoe bat, Rhinolophus capensis. ........................................................................ 21 Figure 2.6 Layout of the bat housing room at the animal housing facility at UCT. .............................. 23 Figure 2.7 Experimental setup in the sound attenuating box (wooden crate, lined on all internal surfaces with sound absorbing foam. Internal dimensions: 77.5 x 38.0 x 38.0 cm), fitted with an ultrasound microphone and an ultrasound speaker, connected to a laptop. The box is monitored by an infrared video camera connected to an external monitor. A small perch is provided on the opposing wall to the speaker. ........................................................................................................ 27 Figure 3.1 Discrimination of echolocation calls by Rhinolophus clivosus. Black indicates the percentage of trials in which the test subjects discriminated between habituation and test playbacks. Grey indicates trials in which test subjects generalised habituation and test playbacks as belonging to the same class of stimuli. Lines with asterisks indicate significant outcomes from post hoc comparisons (Fisher’s exact test, *: P<0.05, **: P< 0.005, ***: P<0.0005). The number of successful trials for each combination is displayed above the bars. ............................................................... 42 iv Figure 3.2 Durations of attentive behaviours during the three analysed experimental phases (HabSTART (light grey), HabEND (no bar) and TEST (dark grey)),), with boxes representing interquartile range and squares representing the median. Whiskers show non-outlier range within 1.5 times the interquartile range. Outliers outside of 1.5 times the interquartile range are presented with circles with extremes (over 2 times) represented by asterisks. The durations of attentive behaviours for the HabEND phase were close to zero and is just visible on this graph. Lines and asterisks indicate outcomes from all possible post hoc comparisons between the TEST phases (*: p < 0.05, **: p < 0.01, ***: p < 0.001). .......................................................................... 43 Figure 3.3 Plot of the first two functions of a DFA on ten acoustic parameters of playback stimuli. Each call of each playback class is plotted (N = 926) and each playback class is colour coded.............. 47 Figure 4.1 Discrimination by Rhinolophus clivosus individuals between playbacks of individually controlled echolocation components. Black indicates the percentage of trials in which the test subjects discriminated between habituation and test playbacks. Grey indicates the percentage of trials in which test subjects generalised habituation and test playbacks as belonging to the same class of stimuli. Lines and asterisks indicate outcomes from all possible post hoc comparisons (Fisher’s exact test, *: P<0.0001). The number of successful trials for each component is displayed above the bars. ............................................................................................................................... 59 Figure 4.2 Durations of attentive behaviours during the three analysed phases (HabSTART (light grey), HabEND (no bar) and TEST (dark grey)), with boxes representing interquartile range. Whiskers show non-outlier range within 1.5 times the interquartile range. Outliers outside of 1.5 times the interquartile range are presented with circles with Extremes (over 2 times) represented by asterisks. The durations of attentive behaviours for the HabEND phase were close to zero and are barely visible on this graph. Lines and asterisks indicate statistical outcomes from all possible post hoc comparisons (*: P<0.01). ......................................................................................................... 61 Figure 4.3 Plot of the first two functions of a DFA on ten acoustic parameters of the male and female playback stimuli. Each call is plotted (N = 1192) and colour coded for gender. ............................ 65 v Figure 4.4 Plot of the first two functions of a DFA on ten acoustic parameters of the high bci and low bci playback stimuli. Each call is plotted (N = 1192) and colour coded for BCI. ............................. 65 Figure 5.1 Experimental setup in the sound attenuating chamber viewed from the top (wooden box, lined on external surfaces with sound absorbing foam (solid lines) or plastic mesh (dashed line)). Internal dimensions: 125 x 125 x 125 cm. Top-Down view), fitted with an two ultrasound speakers, one on either side of a dividing wall broadcasting the playback stimuli simultaneously. The box was monitored by an infrared video camera placed approximately 2 m from the open end and connected to an external monitor. A small holding cage with trap door was positioned on the opposing wall to the speakers, from which the bat was released. ................................................ 80 Figure 5.2 End zone sections set across the open end of the experimental box, viewed from the front. Each side of the dividing wall was subdivided down the middle to create an upper and lower half. The upper half was further subdivided to form a speaker section (a 10 cm perimeter immediately surrounding each speaker) and an upper section (The area above the middle line, excluding the speaker section). Each lower, upper and speaker sections comprised of 25%, 20% and 5% of the total end zone area respectively. ................................................................................................... 82 Figure 5.3 Percentage of landings for each stimulus by Rhinolophus clivosus males. Black indicates the percentage of landings at stimulus 1. Grey indicates the percentage of landings at stimulus 2. .. 88 Figure 5.4 Percentage of landings on either side of the box by Rhinolophus clivosus
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