MALE MATING TACTICS IN THE ROSE BITTERLING (RHODEUS OCELLATUS) AND EUROPEAN BITTERLING (RHODEUS AMARUS) Mara Casalini A Thesis Submitted for the Degree of PhD at the University of St Andrews 2013 Full metadata for this item is available in Research@StAndrews:FullText at: http://research-repository.st-andrews.ac.uk/ Please use this identifier to cite or link to this item: http://hdl.handle.net/10023/3447 This item is protected by original copyright Male mating tactics in the rose bitterling (Rhodeus ocellatus) and European bitterling (Rhodeus amarus) Mara Casalini This thesis is submitted in partial fulfilment for the degree of PhD at the University of St Andrews September 2012 1. Candidate’s declarations I, Mara Casalini, hereby certify that this thesis, which is approximately 40,000 words in length, has been written by me, that it is the record of work carried out by me and that it has not been submitted in any previous application for a higher degree. I was admitted as a research student in July 2008 at the University of Leicester and as a candidate for the degree of PhD in July 2010 at the University of St Andrews; the higher study for which this is a record was carried out in the University of Leicester and of St Andrews between 2008 and 2012. Date………… signature of candidate……………….. 2. Supervisor’s declaration I hereby certify that the candidate has fulfilled the conditions of the Resolution and Regulations appropriate for the degree of PhD in the University of St Andrews and that the candidate is qualified to submit this thesis in application for that degree. Date………… signature of supervisor………………… 3. 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The following is an agreed request by candidate and supervisor regarding the electronic publication of this thesis: Access to printed copy and electronic publication of thesis through the University of St Andrews. Date………… signature of candidate …………… signature of supervisor……… To Matteo. Introduction of the European bitterling D Van Damme et al. Figure 2 Map showing the main areas of intensive carp cul- tivation in Europe around 1500 (modified from Hoffmann 1995). Belgium, they included the basins of the Upper that are found in the region (Throop 1998). One Rhoˆne-Saoˆne, the Rhine-Meuse-Moselle, the Seine- species called ‘pafeduno’ probably refers to the Marne-Loire and the Scheldt-Lys. In central Europe, bitterling, which is still known under this name this industry stretched from central Germany along (‘paffendume’) by fishermen of the Nahe (Portmann the upper Rhine and Main rivers and eastwards 1982; Riethe 1991). The bitterling’s vernacular across the upper basins of the Elbe and Morava in names such as the German ‘paffendume’, the Bohemia and Moravia (present-day Czech Republic). French ‘pe´teuse’ or the Romanian ‘plutica’ are The third centre of production lay in the upper Oder onomatopoeic for the sound that bitterling are and the Vistula basins in Poland. Hoffmann (2000) supposed to produce when captured (authors’ note: cites the following examples of the land surface ‘un pe`t’ is French for a fart). Hildegard describes the claimed by nobles and clergy for fish culture – bitterling as a fish that prefers warm waters, Bohemia (Czech Republic): 25 000 fish ponds; frequenting the shallow nearshore region where it Upper Franconia (south-east Germany): 22 000 fish feeds on small invertebrates and plants (Throop ponds; Upper Silesia (Poland): 25 000 ha; Central 1998). France: 40 000 ha. From the fish ponds in Upper The second oldest reference, and first illustration Silesia 1500 tonnes of live fish (until the late 19th of the European bitterling (Fig. 3), comes from a century freshwater fish were always transferred live Flemish bestiary (Anonymous early 15th century). to market) were transported annually to Crakow This manuscript was produced in Flanders (Bel- (Hoffmann 2002). gium), at that period part of the Burgundian realm Adult bitterling resemble juvenile common carp and share similar habitat preferences. Thus, when young carp ‘seed’ were collected and stored in vats or the flooded compartments of river barges used for transporting live fish it is probable that bitterling were often exported with them. It is notable that all records of the European bitterling in the period approximately 1150–60 are from areas in which carp culture subsequently became important. The earliest bitterling record dates from the mid-12th century and originates from the Rhine-Nahe junc- TheFigure fish 3 The called earliest knownventig representation in the common of the bit- language is named aristosus in Latin because tion near the convent of Bingen (Rinland-Palts, ofterling the (a femalenumerous with an extendedfish bones. ovipositor) It fromis the least palatable of all edible fish. It inhabits south-east Germany). The famous abbess Hildegard anonymous (early 15th century) Flemish bestiary ‘Liber de von Bingen described in her Physica 36 fish taxa freshnaturis rerumwaters creatarum’. as well as those turned brackish by ebb and flood. The meat of the fish is so full of bones that theseÓ 2007 form The Authorsan internal harness, so that it is difficult to chew. 84 Journal compilation Ó 2007In Blackwelladdition, Publishing its Ltd,taste F I S H is and vile F I S H Eand R I E S ,rather8, 79–106 repulsive, which makes it food only suitable for paupers and outcasts. This fish is caught in the following manner: nets are stretched along or across the stream and behind the net an arch-like (hollow) instrument is placed in the water with a little bell suspended in the upper part. When the fish hear the sound of this bell they stupidly rush towards it, being trapped together by the lowered nets. The above proves that this fish possess the sense of hearing. Those fish also like to carry out a kind of group dance, blushing with shame while dancing together. This is something they do (possibly) because it is a way to feed, or because it is a habit of theirs, or because this behaviour is triggered by the presence of others. From the anonymous Flemish bestiary Liber de naturis rerum creatarum (early 15th century). Translation and figure from van Damme et al. 2007. Hence it is the males that fight together and sedulously display their charms before the females. Charles Darwin. From: The Descent of Man and Selection in Relation to Sex (1871). I Acknowledgements First of all I would like to thank my supervisor Doctor Carl Smith. With his support and professional expertise he has greatly contributed to my research experience helping me grow as a researcher and widening my knowledge of the animal world. I would like to thank Professor Martin Reichard of the Institute of Vertebrate Biology, Academy of Sciences of the Czech Republic. He has always heartily welcomed me in Brno, allowed me to use the facilities of his laboratory, and discussed my experimental work and results with insightful comments. A special thank also to Doctor Matej Polačik always ready to help and share his experience and knowledge of bitterling fish. A special thank to Professor S. Tarkan, Doctor Ö. Gayguzus, and Doctor E. Ercan for their help and hospitality in Turkey and for sharing pleasant dinners and after dinners in the guesthouse of Istanbul University in Sapanca. Many thanks to all the PhD students, postdocs and technicians at the Department of Biology of the University of Leicester, where I started my PhD, for their support, advice and friendliness. I am very grateful to all the members of the ‘fish group’ at the University of St Andrews that, during our ‘fish lunches’ at the Gatty, have presented their work- in-progress, discussed results and experimental designs, and given me precious advice for my research. I have learnt a lot from all of them. A special thank to Doctor Miguel Barbosa and Doctor Alfredo Ojanguren: chatting with them about mating systems, mating behaviours, and different ways of life has made cleaning tanks and filters in the aquarium a pleasant job. With Clara, Olivia, Reka, Ian, and Dani I have shared success and worries, coffee breaks, Friday night pub chats, even a hen night and a wedding. A very II hearty thanks to them all. I would like to thank also Muna Agbali: her love for science, her readiness to exchange knowledge and discuss problems, her friendliness and sympathy have made the years of my PhD course a very rich experience. Last, but not least, a very special thank to Matteo. He has patiently listened to endless stories about my bitterlings’ exploits, desperately tried to make me proficient at dealing with advanced computer technology, and, above all, has unconditionally offered me his precious support whenever I needed it. III Table of Contents Abstract .............................................................................................................. 3 List of Figures ...................................................................................................