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Uva-DARE (Digital Academic Repository) UvA-DARE (Digital Academic Repository) Endemism in Sardinia: Evolution, ecology, and conservation in the butterfly Maniola nurag Grill, A. Publication date 2003 Link to publication Citation for published version (APA): Grill, A. (2003). Endemism in Sardinia: Evolution, ecology, and conservation in the butterfly Maniola nurag. IBED, Universiteit van Amsterdam. General rights It is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), other than for strictly personal, individual use, unless the work is under an open content license (like Creative Commons). Disclaimer/Complaints regulations If you believe that digital publication of certain material infringes any of your rights or (privacy) interests, please let the Library know, stating your reasons. In case of a legitimate complaint, the Library will make the material inaccessible and/or remove it from the website. Please Ask the Library: https://uba.uva.nl/en/contact, or a letter to: Library of the University of Amsterdam, Secretariat, Singel 425, 1012 WP Amsterdam, The Netherlands. You will be contacted as soon as possible. UvA-DARE is a service provided by the library of the University of Amsterdam (https://dare.uva.nl) Download date:30 Sep 2021 B^PWMB-a^ip^pMM»!^p^fppq| | differentiationn in the island endemic Maniola wi< < withh Wil van Ginkeï, Gabriel Nève, and StephB,J.M«iken 13$ $ ^-^ ^ Abstract t Inn butterflies, the distribution areas of widespread species and their endemic relativess are usually vicariant. In Sardinia, the ranges of an endemic and a widespreadd Maniola species overlap, and the two species possibly hybridise. In this paper,, we analyse patterns of genetic differentiation in Maniola nurag and Maniola jurtinajurtina from Sardinia by means of allozyme markers, compare them to mainland M.M. jurtina populations, and interpret the data with regard to the endemic species' evolutionaryy history. Sardinian M. nurag and M. jurtina have equally high levels off genetic variation (H = 0.141-0.270; H = 0.137-0.189) as mainland M. jurtina (H(Hnainlandnainland== 0.141-0.236). Total genie diversity at fifteen polymorphic loci is mostly duee to within population variation (FIS). The close relationship of the two species is illustratedd by the fact, that 63 of the 76 alleles screened are shared by both species. Smalll genetic distance between them (Nei's D = 0.21) indicates that divergence initiatedd after the desiccation of the Mediterranean (+ 3 ma ago), and was possibly associatedd with the abrupt climate changes at the turn from Pliocene to Pleistocene (1.8-33 ma). Geographic patterns in allozyme allele frequencies hint at the existence off hybrizymes, and suggest the presence of hybrids in areas where M. nurag and M.M. jurtina are sympatric. Island populations of neither species show signs of loss off genetic diversity, inbreeding, or bottlenecks. We propose that M. nurag did not resultt from vicariance or dispersal, but originated under sympatric or parapatric conditions,, as a consequence of local adaptation along an environmental gradient. Keywords:: Maniola nurag, butterfly, genetic population structure, hybridisation, endemic,, allozymes, inbreeding, Lepidoptera, Nymphalidae, divergence time Introduction n AA species is usually genetically structured over space and in time. Historical founderr events, bottlenecks, and gene-flow are important evolutionary agents responsiblee for changes in the genetical structure of populations. Assessing genetic variationn across geographic areas can thus provide means to trace the history of thesee populations and eventually the history of species (Avise, 1994; Hewitt, 1999; Schmitt&Seitz,2002). Severall speciation models have been proposed, which are habitually characterized byy the level of gene-flow between diverging populations during the initial stages off speciation {Dobzhansky, 1940). Population divergence in the presence of gene- floww was often considered to be unrealistic. However, a number of theoretical 136 6 studiess have reported the plausibility of sympatric and parapatric speciation, and havee shown that spatially localized interactions along environmental gradients cann facilitate species' differentiation (e.g., Kondrashov & Kondrashov, 1999; Doebelii & Dieckmann, 2000; 2003). Despite this growing theoretical evidence that ecologicallyy driven speciation can occur, empirical studies showing examples for suchh speciation modes still remain scarce (Ogden & Thorpe, 2002; Scriber, 2002; Lushaii et al., 2003). Earlier sympatric speciation models involved ecologically drivenn reproductive isolation associated with adaptation to alternative resources (nichee shift), as was elegantly shown for the host races in the tephritid fly Rhagoletis pomonellapomonella (Bush, 1994) or the large cactus finch Geospiza consirostris (Grant & Grant, 1989).. Recent modelling advances suggest that competition for continuously distributedd resources, driven by sexual selection against intermediate phenotypes, couldd be the driving force for sympatric speciation (Doebeli & Dieckmann, 2003). Intermediatee phenotypes procure fewer resources as a consequence of density- andd frequency-dependent selection, and are selected against under disruptive selectionn (Turelli et al., 2001). Hybridd zones form an ideal environment to study sympatric speciation (Arnold, 1997).. A 'hybrid zone' sensn Arnold (1997) is a geographical area where "two populationss of individuals that are distinguishable on the basis of one or more heritablee characters overlap spatially and temporally and cross to form viable and att least partially fertile offspring." In such a parapatric situation, gene-flow can sloww down or even inhibit differentiation by spreading favourable alleles across the hybridd zone (Kim & Rieseberg, 1999), whereas reinforcement can cause prezygotic reproductivee isolation (Turelli et al., 2001). Reinforcement intensifies mate preferencee (Dobzhansky & Pavlovsky, 1957; Butlin, 1995), and can lead to character displacement.. With character displacement, the differences between sympatric populationss of two species are accentuated as a result of reproductive or ecological interactionss between them (Futuyma, 1998). Although character displacement has generallyy been interpreted as an evolutionary response to secondary contact, it cann also evolve in situ across an environmental gradient, despite continuing gene- floww (Turelli et al., 2001). The existence of hybrid zones and steep genetic clines (Schilthuizenn et al, 1999, Lushai et al., 2003) shows that selection can dominate gene-floww over small spatial scales and therefore allow for parapatric divergence. Inn many hybrid zones, particular allozymes called 'hybrizymes' (Woodruff, 1989) havee been found, representing alleles that are not present or very rare in the parentall taxa, and reflect novel genetic variation (Schilthuizen & Gittenberger, 1994b;; Arntzen, 2001). Hybrid zones have been extensively investigated in plants, andd also in animals (e.g. Barton & Hewitt, 1985; Hewitt, 1988,1999; Schilthuizen & 137 7 Lombaerts/1995;; Arntzen, 2001; Capula, 2002) but only rarely so in Lepidoptera (Aagaard,, 2002; Scriber, 2002; Lushai et al, 2003). Thee distribution areas of widespread species and their endemic relatives are usually disjunctt in butterflies (Dennis et al., 2000). In the genus Maniola (Lepidoptera, Nymphalidae),, however, the Sardinian endemic Maniola nurag (GHILIANI 1852) andd its widespread close relative, Maniola jurtina (L. 1758), are found in sympatry andd possibly hybridise (Grill et al, 2003b, 2003d). In order to find out whether thee present sympatric occurrence of the two species can be best explained under thee assumption of a sympatric, parapatric, or allopatric mode of speciation, we investigatee the population genetic structure in a number of island populations of bothh species, and compare these to continental populations of M. jurtina by means off allozyme markers. As we found ecological as well as morphological support suggestingg that M. nurag and Al jurtina possibly hybridise in Sardinia (Grill et al.,al., 2003b; 2003c), we further evaluate the probability of hybrid occurrence in the Sardiniann Maniola. Allozymess are co-dominant markers and efficient to study population differentiationn in Lepidoptera because of the large number of polymorphic loci {e.g., Raijmannn & Menken, 2000; Nève, 2000; Schmitt & Seitz, 2002), and also provide uss with a straightforward tool to detect interspecific hybridisation (Menken & Ulenber,, 1987; Schilthuizen & Gittenberger, 1994b; Schilthuizen & Lombaerts 1995; Arntzen,, 2001; Capula 2002); diagnostic loci differentiate between species (Hewitt, 1988;; Grant & Grant, 1996; Schilthuizen et al., 1999), and consequently can reveal whetherr hybridisation takes place. Iff the present sympatric occurrence of M. nurag and Al jurtina in Sardinia resulted fromm a sympatric or parapatric speciation event we would expect to find evidence forr reinforcement or disruptive selection on traits that are associated with the use off alternative niches (Mayr, 1963, Bush, 1969). In an early phase of differentiation, mostt alleles at polymorphic loci are still shared between the populations in similarr frequencies, and gene-flow between the diverging populations is large. Geneticc regions that are involved in differential adaptation, however, continue too diverge through selection. In later stages of differentiation, gene-flow will be furtherr reduced, and neutral
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