Vision in echolocating bats - Fladdermus.net

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multimodality (Pettigrew et al. 1983; Schmidt 1988; Hessel & Schmidt 1994). In a two choice test, two phyllostomid bats (Desmodus rotundus and Phyllostomus discolor) were trained to respond to a combination of visual, olfactory and acoustic stimuli, and were then presented with one of the three modalities separately (Schmidt et al. 1988). It was found that Phyllostomus discolor chose the visual stimuli to a higher degree, whereas Desmodus rotundus preferred the passive acoustic stimuli. However, both bats were able to respond to all three modalities, although responses to the olfactory stimuli needed additional training, as also noted by Hessel and Schmidt (1994), when studying Carollia perspicillata. However when the Carollia had learned to respond to the olfactory cue, this became the preferred stimuli, which was not the case with Desmodus or Phyllostomus, which both used two other senses. This clearly shows that bats use an array of different senses in the field, and that ecology, feeding strategies and behavioural context all influence the use of different modalities. Echolocation may be the most important innovation throughout bat evolution, allowing these animals to explore a niche of their own. But there is more to the sensory ecology of microchiropteran bats, where vision is an important piece of the puzzle and certainly needs further attention in the future. 37
Acknowledgements First of all I would like to acknowledge all the co-authors of this thesis for obvious reasons, and I wish to thank Olof Helje for making the splendid bat illustrations. Then I wish to thank my supervisor and mentor -Jens Rydell – and also, many thanks to the rest of the Rydell family for your great hospitality. There are several people having answered several more or less stupid questions on bats, vision, statistics, experimental design, the meaning of life and other various topics throughout the years. Especially I would like to thank Donald R Griffin and Brock Fenton, but also Eric Warrant, Dan E Nilsson, Tom, Gareth Jones, Susan Swift, Kristina Mieziewska, Winston Lancaster; and of course, many thanks to Cajsa, John G, Christoffer, Gim, Jen, Christin, John R, Marc, Jenny, Kalle, Kristina, Cess, Bomull, Tobias, Annika, Christoffer and Staffan, just to mention a few of you. Just as many people have helped me to make the every day work possible; at the department, by joining me in the field, on conferences, courses and work shops and to some extent even in the lab (although some may think I do not know what that is); having helped me arrange field work and experiments, being guides, eye suppliers, hosts, or just good company during batting. Especially I wish to thank Monica, Tompa and Cajsa, thanks also to Maria, Héctor, Henke, Luis-Bernardo, Hans, Marie, Gabriela, Stefan, Åsa, Per, Britt-Louise, Anna, Lee, Dr. F-Jo, Blomman, Karl-Johan, Stefan, Sean, Jorge, Jenny, Andreas, Eric, Lars-Erik, Cess, Annemarie, Magnus, Berndt, Dave, Anne-Sofie, Bengt, Mia, Lilioth, the “NASBR and Chamela-students”, the Lövhaugs, all other department employees not mentioned, and of course Mexican hospitality and British humour. For unknown reasons, I have been deeply involved, not only in research and teaching, but also in the work of the faculty board, the Swedish Association of Scientists and the Students’ Union. I wish to thank the various members of all the different working groups and committees, not at least Cajsa, Marie, Stefan Henrik, and Andreas. There are other things but science, like having almost normal conversations, sharing stupid ideas, trying to do music, lying on beaches, e-mailing, having coffee and drinking beer, and presumably some other stuff as well. For those things, I wish to thank Per and Andreas for helping me to create “PSL” which for a while brought order to my life, much in the same way as “Johnny” did, only different. I wish to thank the e-mailers, the floorballers, the chatters, the Herb Boys crew and fan club, the coffee drinkers and the travellers. There is no doubt that the Friday after work sessions have been almost as important as the actual research for being able to finish this thesis. I wish to acknowledge the most frequent ones: Christoffer, Anna, Viktoria, Ågot and lately Jenny P. But of course, Sara, Erik, Fredrik, Jenny T, Linda, Tove, Goran and a whole bunch of other youknow-who-you-are. There are two persons having shared my biology- as well as my non biology-time, to a larger extent than perhaps any others: first of all, thank you Cajsa for not bringing your calculator; and for numerous moments thereafter, and second, Tompa, for making everyday April fools day. Thank you all on the second floor: Per, Bengt, Stig, Gunnar, Anders, Inger, Lena, Åke, Jan, Sebbe, Björn, Jenny, Malin, Urban, Anna H, Anna Z, Monica, Susanne, Ulla, Mare, Christoffer, Arne, Marcus, other hangarounds, not mentioned, past and present. Thank you mom, dad, and Kristian, and thanks to all friends in the real world. My work has been funded by Kungliga och Hvitfeldtska Stiftelsen, Lunds Djurskyddsfond, Knut och Alice Wallenbergs Stiftelse, Folke Eklöf, Adlerbertska Forskningsstiftelsen, Wilhelm och Martina Lundgrens Vetenskapsfond, Kungliga vetenskaps- och Vitterhetssamhället i Göteborg, Stiftelserna Paul och Marie Berghaus Donationsfond, J A Ahlstrands Testamentsfond, Lars Hiertas Minne, and Rådman & Fru Ernst Collianders Stiftelse FVÄ, and of course CSN. Finally I wish to acknowledge (please fill out your name); believe me you are not really forgotten. 38
Page 1 and 2: Vision in echolocating bats Johan E
Page 3: A doctoral thesis at a university i
Page 6 and 7: to be precedence of vision over son
Page 8 and 9: och för att undvika hinder på vä
Page 11 and 12: B ats (Order: Chiroptera) are among
Page 13 and 14: VISION IN ECHOLOCATING BATS The mic
Page 15 and 16: The microchiropteran eyes are shape
Page 17 and 18: structures, and the dorsal lateral
Page 19 and 20: 10 lux, which is equivalent to the
Page 21 and 22: The device used for the optomotor r
Page 23 and 24: Tab 2 - cont Light Visual Species (
Page 25 and 26: jamaicensis, Desmodus rotundus and
Page 27 and 28: (Davis & Barbour 1965) or when they
Page 29 and 30: Insectivores and carnivores For bat
Page 31 and 32: Moreover, the eyes of Macrotus cali
Page 33 and 34: combined with broadband echolocatio
Page 35 and 36: Multimodality - vision and echoloca
Page 37: superior colliculus. It has been sh
Page 41 and 42: Buchler, E. R. & Wasilewski, P. J.
Page 43 and 44: Heffner, R. S. 1997. Comparative st
Page 45 and 46: Masterson, F. A. & Ellins, S. R. 19
Page 47: Suthers, R. A. & Chase, J. 1966. Vi
Page 51 and 52: Animal Behaviour - In Press Use of
Page 53 and 54: the ability of brown long eared bat
Page 55 and 56: General observations RESULTS When r
Page 57 and 58: are influenced to some extent by th
Page 59 and 60: Eklöf, J., Tranefors, T. & Vazquez
Page 61 and 62: Table 1. - The number of feeding at
Page 63: “The Eocene brought mammals mean
Page 73 and 74: Submitted manuscript Vision complem
Page 75 and 76: (dorsal) side up and the other had
Page 77 and 78: (Anderson et al. 1998). Hence, the
Page 79: Figure 1. - Frequency of attacks by
Page 83 and 84: Manuscript Visual acuity and eye si
Page 85 and 86: computer screen. The bats were rele
Page 87 and 88: study. On overcast nights the amoun
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Koay, G., Kearns, D., Heffner, H. E
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Tab. 1. - Eye diameter and optomoto
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“The bat that flits at close of E
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