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Charles University Faculty of Science Department of Zoology Charles University Faculty of Science Department of Zoology Suckling and allosuckling in ungulates RNDr. Jan Pluháček, Ph.D. Habilitation Thesis Praha 2017 1 Motto: Mnoho dat z biologie jednotlivých druhů jako je délka březosti, doba kojení či vývoj mláďat, je možno získat jedině v zoologických zahradách. (A lot of data on biology of individual species as gravidity, time spent by lactation or the development of offspring could be received only in the zoos.) Zdeněk Veselovský 2 Suckling and allosuckling in Grévy’s zebra (Equus grevyi) at Dvůr Králové Zoo in summer 2010 Suckling in common hippopotamus (Hippopotamus amphibius) in Ostrava Zoo in spring 2011 3 Content Content ....................................................................................................................................... 4 List of publications included into this thesis .............................................................................. 5 Prologue ..................................................................................................................................... 6 Acknowledgements .................................................................................................................... 7 Introduction or why suckling behaviour and why zebras? ......................................................... 7 On importance of suckling behaviour ........................................................................................ 9 The mismeasure of suckling behaviour .................................................................................... 11 At the Old Giraffe House in Dvůr Králové Zoo ....................................................................... 13 On the magic sex ...................................................................................................................... 15 Striped horse or striped ass ....................................................................................................... 18 From the left to the right or the laterality of suckling .............................................................. 21 Allosuckling as the extreme investment ................................................................................... 23 On allosuckling of mysterious river horse and elegant beauty that forgot to count ................. 26 Epilogue ................................................................................................................................... 30 References ................................................................................................................................ 31 4 List of publications included into this thesis Pluháček, J., Bartoš, L., Doležalová, M., & Bartošová-Víchová, J. (2007). Sex of the foetus determines the time of weaning of the previous offspring of captive plains zebra (Equus burchelli). Applied Animal Behaviour Science, 105, 192-204. Pluháček, J., Bartoš, L., & Bartošová, J. (2010). Mother-offspring conflict in captive plains zebra (Equus burchellii): suckling bout duration. Applied Animal Behaviour Science, 122 (2-4), 127-132. Pluháček, J., Bartoš, L., Bartošová, J., & Kotrba, R. (2010). Feeding behaviour affects nursing behaviour in captive plains zebra (Equus burchellii). Applied Animal Behaviour Science, 128, 97-102. Pluháček, J., Bartošová, J., & Bartoš, L. (2010). Suckling behavior in captive plains zebra (Equus burchellii): sex differences in foal behavior. Journal of Animal Science, 88 (1), 131-136. Pluháček, J., & Bartošová, J. (2011). A case of suckling and allosuckling behaviour in captive common hippopotamus. Mammalian Biology, 76, 380-383. Pluháček, J., Bartošová, J., & Bartoš, L. (2011). A case of adoption and allonursing in captive plains zebra (Equus burchellii). Behavioural Processes, 86, 174-177. Pluháček, J., Bartošová, J., & Bartoš, L. (2011). Further evidence for sex differences in suckling behaviour of captive plains zebra foals. Acta Ethologica, 14, 91-95. Olléová, M., Pluháček, J., & King, S. R. B. (2012). Effect of social system on allosuckling and adoption in zebras. Journal of Zoology, 288 (2), 127-134. Pluháček, J., Olléová, M., Bartošová, J., & Bartoš, L. (2012). Effect of ecological adaptation on suckling behaviour in three zebra species. Behaviour, 149(13-14), 1395-1411. Pluháček, J., Olléová, M., Bartošová, J., Pluháčková, J., & Bartoš, L. (2013). Laterality of suckling behaviour in three zebra species. Laterality: Asymmetries of Body, Brain and Cognition, 18 (3), 349-364. Pluháček, J., Olléová, M., Bartoš, L., & Bartošová, J. (2014). Time spent suckling is affected by different social organization in three zebra species. Journal of Zoology, 292 (1), 10- 17. Gloneková, M., Brandlová, K., & Pluháček, J. (2016). Stealing milk by young and reciprocal mothers: high incidence of allonursing in giraffes, Giraffa camelopardalis. Animal Behaviour, 113(1), 113-123. 5 Prologue Saturday, January, the 9th, 1999 was quite a typical day in the winter season in central Europe. The ground was covered with the remnants of snow and the air temperature was a little higher above zero. In the heated “Old Giraffe House” at Dvůr Králové nad Labem Zoo I started my observation of suckling behaviour of plains zebra. At 8:00:38 I recorded the first suckling bout. It lasted 51 seconds. It was performed by “Irbil”, son of “Slávek” and “Irkasa” in the herd of maneless zebra (Equus quagga borensis). Since then I recorded other 4815 suckling bouts and 4297 suckling attempts in captive zebra during the following 12 years. Analyses of that data as well as that recorded by my students resulted in 10 scientific publications which form the main content of this thesis. In addition, the thesis includes also two papers on suckling behaviour of two different ungulates: common hippopotamus (Hippopotamus amphibius) and giraffe (Giraffa camelopardalis). All these papers are based on data collected in Czech zoos. I would like to stress that ethological studies in the zoos were not favoured in Czechia/Czechoslovakia in the past. Moreover, I faced from time to time critical comments by reviewers of scientific journals that the data coming from captivity are not worthy to be published. The most common comment is that the captivity modified animal behaviour too highly. I would like to note that I did not hear such a comment over any study using data on rodents kept in laboratories or on fish in aquaria, although it could be the absence of my knowledge. I do not want to say that captivity does not modify animal behaviour. It does for sure. Nevertheless, I would like to say that it is better to study any phenomenon in captive animals than to resign to study it at all (Calisi & Bentley 2009). Moreover, the observation of zoo animals involves several advantages as we are well informed on the history of all individuals including their exact age, reproductive history or kinship. I think that among the best examples of an animal which would be worthy to be studied in captivity is hippopotamus. To observe hippopotamuses in the wild might be quite a dangerous activity. The observed individual may easily disappear under the water full of crocodiles. It rejects all possible marking and it is able to wash the colour marking out quickly. When using telemetry, the hippopotamus destroys the collar in the water and mud environment. Last but not least, the longevity of hippopotamus is higher than that of many grant agencies. My interest in animals was based on my frequent visits of zoos in my childhood. Later, I received nice inspiration by reading books written by prof. Zdeněk Veselovský, Dr 6 Luděk Dobroruka and Dr Jiří Volf (Czechoslovak researchers who worked at zoos) who supported the ethological research at zoos and considered research as one of the main role of modern zoos. I hope that the publications included in this thesis could provide at least some evidence of this role. Acknowledgements I would like to express my biggest thanks to Luděk Bartoš who taught me all methods in ethology, helped me a lot to analyse the data, commented the manuscripts valuably, and especially supported me during the whole time of observation of zebras at various zoos. My thanks are due to all co-authors of the publications included in this thesis, namely Jitka Bartošová, Sarah R. B. King, Markéta Gloneková, Radim Kotrba, Karolína Brandlová, Miroslava Doležalová, and one of my first students Michaela Levá (Michaela Olléová during the time of observation). I would like to gratefully acknowledge the assistance of the staff of the Zoological Garden at Dvůr Králové and Ostrava, in particular Luděk Čulík, Markéta Čulíková, Aleš Kopecký, Jiří Soumar, Miroslava Kubelková, Pavel Moucha, Radek Hlávka, Barbara Raková, Kristina Tomášová, Miroslav Špráchal, Jan Ždárek, Václav Hák, Martin Vyhledal, Dagmar Janková, Jiří Hrubý, Eva Zajoncová, Robert Kopia, Rostislav Střižík, Karin Skýbová, Lenka Málcová, Anna Leštínská, Liběna Hájková and Petr Čolas. I am indebted to my father Alois Pluháček for technical help. The scientific articles we published were highly improved by valuable comments from my very kind colleagues from the Department of Ethology, Institute of Animal Science, namely Radka Šárová, Francisco Ceacero, Martina Komárková, Vratislav Kšáda, Petr Janovský, and Marek Špinka. Part of this work was supported by grant no. 523/08/P313 from the Czech Science Foundation. Last, I would like to thank my son Ondřej for his patience during writing this thesis. Introduction or why suckling behaviour and why zebras? I would like to start this text by
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