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Galerucella Nymphaeae PDF hosted at the Radboud Repository of the Radboud University Nijmegen The following full text is a publisher's version. For additional information about this publication click this link. http://hdl.handle.net/2066/19064 Please be advised that this information was generated on 2021-10-08 and may be subject to change. Evolution in action: Host race formation in Galerucella nymphaeae Stephanie Pappers 6WHOOLQJHQ behorende bij het proefschrift Evolution in action: host race formation in Galerucella nymphaeae Stephanie Pappers 1. Het waterleliehaantje, Galerucella nymphaeae, bestaat uit minstens twee gastheerrassen. 2. Differentiatie is mogelijk zonder geografische barrière, ook in de natuur. 3. Nothing in biology makes sense except in the light of evolution. 'RE]KDQVN\ $PHULFDQ %LRORJ\ 7HDFKHU 4. Als de vroege christenen beter naar Empedocles in plaats van Aristoteles hadden geluisterd had Darwin het niet zo moeilijk gehad. 5. De biologie heeft mij over de Schepper geleerd dat Hij in ieder geval een bijzondere voorkeur voor kevers heeft. 'H %ULWVH JHQHWLFXV -%6 +DOGDQH YROJHQV +XWFKLQVRQ $PHULFDQ 1DWXUDOLVW 6. Met de toenemende vergrijzing bij de universiteit zal ook het papierverbruik toenemen, al is het alleen al omdat een steeds groter lettertype nodig is. 7. Aangezien de meeste stadsduiven afstammen van verwilderde uitheemse rotsduiven zouden ze dus, alleen al uit oogpunt van natuurbescherming en het tegengaan van genetische vervuiling van de inheemse duiven- soorten, bestreden moeten worden. 8. Natuur is overal waar je mobieltje het niet doet. 0HGHZHUNHU 1DWXXUPRQXPHQWHQ DDQJHKDDOG LQ 'H 9RONVNUDQW DSULO 9. Wie aardige mensen wil ontmoeten kan het beste bloeddonor worden. 10. Treinreizen bevordert literatuurkennis. Evolution in action: Host race formation in Galerucella nymphaeae een wetenschappelijke proeve op het gebied van de Natuurwetenschappen, Wiskunde en Informatica Proefschrift ter verkrijging van de graad van doctor aan de Katholieke Universiteit Nijmegen, volgens besluit van het College van Decanen in het openbaar te verdedigen op maandag 17 december 2001 des namiddags om 3.30 uur precies door Stephanie Maria Pappers geboren op 3 oktober 1972 te Schiedam Promotor: Prof. Dr. J.M. van Groenendael Co-promotores: Dr. N.J. Ouborg Dr. G. van der Velde Manuscriptcommissie: Prof. Dr. J.J.M. van Alphen (Universiteit van Leiden) Prof. Dr. S.B.J. Menken (Universiteit van Amsterdam) Prof. Dr. L.E.M. Vet (NIOO-KNAW) Omslag: collage van gefotografeerde eipakketten van Galerucella nymphaeae door Marij Orbons Omslag ontwerp: Jolanda Hiddink, Afdeling Grafische Vormgeving, KUN ISBN: 90-9015225-3 © 2001 S.M. Pappers Parts of this material are allowed to be reproduced or utilised as long as their source is mentioned. Voor mijn ouders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entral to biology is the theory of evolution, the adaptive change between gen- erations within a population of a species. Most evolutionary biologists tend to believe that the history of evolutionary biology begins with Darwin. However, ancient Greek philosophers already thought about evolution. Anaximander (611?-547? BC) taught his students that life arose in water, complex forms of life arose from simpler forms and that humans arose from fish that left the seas to live on dry land. Empedocles (495?-435? BC) added an explanation of the evolu- tionary process to the ideas of Anaximander, namely that only organisms with harmonious combination of body parts, which were brought together by forces of attraction, survived. Furthermore, he stated that the harmonious combina- tions of structures arose by chance, after a number of attempts. Thus, his ideas already contains the concept of adaptation. Later, these ideas were replaced by the ideas of the Classical School, to which Plato (427?-347 BC) and Aristotle (348- 322 BC) belonged. They emphasised the final, perfection in organisms, thus re- jecting the idea of change (Störig 1996). This idea of ‘fixity of species’ was com- monly accepted by scientists until the beginning of the 19th century, when La- marck (1744-1829) published his theory of evolution. And it took until 1859 be- fore Darwin published his On the origin of species by means of natural selection (1859). The observation of variation in the finches of the Galapagos islands and other birds in South America during his trip with the Beagle (1832-1837) led Darwin to the idea that species can change. By that time, however, he had no idea of what caused such change. In his autobiography, he wrote the following explanation for why species change: “In October 1838, that is fifteen months after I had begun my systematic enquiry, I happened to read for amusement ‘Malthus on population’, and being well prepared to appreciate the struggle for existence which every- where goes on from long-continued observation of the habits of animals and plants, it at once struck me that under these circumstances favourable variations would tend to be preserved and unfavourable ones to be de- stroyed. The results of this would be the formation of a new species.” (cited from Ridley 1996). Thus, Darwin raised the idea of how species could be formed in 1859, however, it took until 1939, for the first symposium on ’speciation’ took place (Cole 1940). Since then, the term ‘speciation’ is commonly used by all evolutionary biologists. One of the difficulties in speciation research was, and still is, the problem of a useful definition of a species. So far, many species concepts have been proposed. The typological species concept is probably the oldest, since it originated with Plato and Aristotle. According to this concept, a species represents some ideal &KDSWHU form, of which individual variation is merely the imperfect expression. Closely related to this concept is the morphological species definition, which states that “organisms of a certain species are more similar to each other than to organisms of other species”. These definitions, however, do not take into account the possi- bility of convergent evolution and neglects what we now call ‘morphologically indistinguishable cryptic sister species’. Furthermore, according to these two species concepts, species are static: they cannot change over time or space. Nowadays, the most widely used concept, which is also easy to understand, is the biological species concept (Dobzhansky 1937, Mayr 1963). This concept de- fines a species as “a group of interbreeding or potentially interbreeding popula- tions with fertile and viable offspring and incapable of breeding with other such populations”. Of course, this concept gives many problems in asexually repro- ducing organisms, but in most sexually reproducing organisms it is a testable definition. Other species concepts, such as the phylogenetic or evolutionary spe- cies concepts may be theoretically more valid than the biological species con- cept, but they hardly can be tested empirically. For instance, the evolutionary species concept defines a species as a single lineage of ancestor-descendent populations which maintains its identity from other such lineages and which has its own evolutionary tendencies and historical fate (Wiley 1981), but the lat- ter part of this definition is difficult to test or prove. Therefore, the biological species concept will be used in this thesis. Like the view on the definition of a species has changed over time, so did the view on the causes of speciation. The crucial event, for the origin of a new spe- cies, is reproductive isolation. Theories about speciation differ in how such isola- tion is achieved. In the classical speciation model, reproductive isolation is caused by some extrinsic barrier. By such a barrier, widely distributed species become subdivided into two or more relatively large populations. After the bar- rier has interrupted gene flow, genetic differences begin to accumulate between the two or more daughter populations as a result of random genetic drift and different selection regimes at each side of the barrier. Mayr (1942) called this process allopatric speciation (Greek: allos=other, patria=homeland). Among the numerous examples of allopatric speciation, is of course the classical example of the finches of the Galapagos islands. Darwin (1859) studied 13 different species of finches which are found on different islands in the Galapagos chain and no- where else on earth. Darwin believed that all species have a small group of common ancestors who came to the Galapagos from the South American mainland by a rare event. Among other traits, the species differ in beak mor- phology, which is adapted to feed on different types of seeds or insects. These birds have provided a case study of how a single species
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