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Genetic and Ecological Differentiation Between the Butterfly Sisterspecies

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Genetic and Ecological Differentiation Between the Butterfly Sisterspecies Contributions to Zoology, 71 (4) I3I-I39 (2002) SPB Academic Publishing bv, The Hague Genetic and ecological differentiation between the butterfly sisterspecies Colias alfacariensis and Colias hyale Daniel+F.R. Cleary Henri Descimon² and Menken¹ ¹, Steph+B.J. 1 Institute for Biodiversity and Ecosystem Dynamics (IBED), Faculty ofScience, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands, [email protected]; 2 Laboratoire de Systematique Evolutive, Universite deProvence, Marseille, France Keywords: Lepidoptera, Pieridae, allozymes, population structure, gene flow Abstract Habitat preference 134 Population analysis 134 Discussion 136 Clear habitat separation between the sister species Colias alfa- Acknowledgements 138 cariensis and C. hyale is shown when occurring sympatrically. References 138 Colias hyale is found more often in moist cultivated pastures while Colias alfacariensis is more abundant in dry uncultivated habitat. Out of a total of 16 loci, no diagnostic loci were found between C. alfacariensis and C. hyale, and both species shared Introduction most major polymorphisms. Exceptions were the marked dif- ferences in allele frequencies at the HK locus and only C, hyale, Coexistence of species requires the existence of but not C. alfacariensis was further invariable at the GOT2 lo- evolutionarily continual trade-offs in cus, which is usually highly polymorphic in the Pieridae. Colias interspecific the of hyale has a significantly lower level of heterozygosity than its abilities species to deal with factors that limit sister species C. alfacariensis. In Colias alfacariensis hetero- their fitness and abundance. These include trade- zygosity is highest in the Alps and lowest in the low-lying region offs between competitive abilities and dispersal ofNorthern France, Both show levels of flow species high gene abilities, susceptibility to disease, herbivory or pre- over a large geographic area. Within C. alfacariensis, but not dation, between abilities to live off condi- in C. hyale, the F value ofthe PGI locus is significantly diffe- average ST rent tions or abilities to resource and from zero effectively separating the species into populations exploit pulses, ’ with high levels ofthe ’ b allele to the west and North, and low between abilities to compete for alternative resources levels of the allele in the Alps and Italy. This could point to in a heterogeneous landscape (Tilman, 2000). Feed- selection within the PGI locus in line with the well established ing ecology, reproductive behaviour, and physiol- pattern ofselection at the PGI locus in other species ofColias. ogy are characteristics of a species niche. Niches Glaciations have been an important force in shaping the evolu- tend to be scales tionary history ofEuropeanbiota, leading to extinction, but also conservative over time associated evolve into the allowing new species to newly available land as with sister taxon pairs in, for example, birds, mam- the ice sheets retreated. The and distributional genetic pattern mals, and butterflies but this conservatism breaks found between both Colias species suggests that habitat shifts down at the level of families (Peterson et ah, 1999). and subsequentadaptation during glaciations could have played By analysing the present day difference in genetic an important role in their speciation. makeup and the often subtle differences in ecol- within ogy between species a pair of sister species we can an idea of the forces that Contents get evolutionary led to their differentiation into separate species. The two Introduction 131 present study concerns species of the Material and methods 132 genus Colias [Lepidoptera: Pieridae] namely C. Habitat preference 132„ alfacariensis and C. hyale (Brunton, 1998). Colias Population analysis 132 is one of the most widespread butterfly genera in Results 134 It the world. is present on all continents except Downloaded from Brill.com10/03/2021 08:23:00PM via free access - 132 D.F.R. Cleary et al. Genetic and ecological differntiation between butterfly sisterspecies and from the Antarctic Colias Australia, ranges (e.g., Materials and methods the Arctic antarctica) to (e.g., Colias boothi), across Asia, Africa, North and South America, from the Habitat preference lowlands to the highest mountains. Colias and C. sister alfacariensis hyale are spe- Individuals of both C. alfacariensis and C. hyale cies (Brunton, 1998), and are difficult to tell apart were sampled for an allozyme study in France and due to the large variation in wing patterns and other Italy in the summer of 1992. Species determina- characters of the butterflies. morphological In gen- tion was based on the studies of Berger and Fontaine eral, C. hyale has a more northerly and C. alfaca- (1947), Jarvis (1955), and Dutreix (1980). Habitat riensis a more southerly distribution with a large preference between C. alfacariensis and C. hyale of in In region overlap central Europe. recent years was studied by means of field observations in the C. has become in northern hyale increasingly rare French Alps where both species occur sympatri- clue in Europe to abandonment agricultural areas cally. of its larval food plant Medicago sativa (Descimon, A total of 920 adult C. alfacariensis and 1155 C. th (1994(5)). adult hyale were observed from the 14 of Au- th Colias alfacariensis flies in May-June and Au- gust until the 8 of September, 1974 by HD. The gust-September in two, and in southern Europe location and geographical position of all sites is sometimes in three generations. Food plants are presented in Table 1. During the field observations and Coronilla varia. the environmental Hippocrepis comosa Colias following variables were noted hyale has similar flight periods and the food plants at each locality: 1. Aspect, 2. Vegetation (cultivated Vida Coronilla - are Medicago sativa, cracca, spp., uncultivated), 3. Humidity (dry - irrigated), 4. Lotus and other Altitude - spp., Cytisus spp., Trifolium spp., 5. Date, 6. Weather (warm cold), in ad- Fabaceae dition the (Higgins and Riley, 1970). Both species to frequency of occurrence of adults of arc thus oligophagous on Fabaceae although C. hyale C. alfacariensis and C. hyale. has of food a larger range potential plants. The relative abundances of both species were The of this is the the goal study to analyse ecology regressed onto first axis of a principal compo- of both based and genetical population structure species nents analysis (PCA on the environmental and to to into the differentiation try gain insight variables and using Statistica for Windows, ver- process that produced both present-day taxa. Lepi- sion 5, StatSoft Inc.) to see if both species varied for doptera are an important source understanding significantly (positively or negatively) in their re- the genetics of speciation (Menken et ah, 1992; sponse to this synthetic environmental gradient. The Ritchie and Phillips, 1998). The analysis of spatial relative abundance here is the absolute abundance pattern (e.g., relative densities and habitat prefer- of species A at a given site divided by the absolute Mark- ences), direct measures of migration (e.g., abundances of both species at the same site. We abundance here rclcase-rccapture), and indirect measures (e.g.fi use relative because of the high all for under- in genetic differentiation) are important variance the absolute abundance among sites. standing how populations are structured. Lewis et al. (1997; and refs therein) suggest that at least two of methods should be in order these applied to pro- Population analysis vide a good indication of population structure. The we address are the fol- specific questions For the allozyme electrophoresis analyses, popu- Do both differ in their habitat of both lowing. 1) species lations Colias species were sampled in the What is the preference? 2) spatial, genetical popu- French Alps and other localities, including the Pro- lation structure of both species? vence, Northern France and Italy (Table 1). After capture the butterflies were stored in liquid ni- head trogen. The and thorax were subsequently in in 180 ml of M homogenised the laboratory a 0.1 Tris-HCl buffer (pH 8.0) and the extracts were cen- Downloaded from Brill.com10/03/2021 08:23:00PM via free access Contributions to Zoology, 71 (4) - 2002 133 Table I. Location and of all sites. geographical position sampling Allozyme indicates that the population sampled was used in allozyme electrophoretic analysis. Field indicates that the site was used in the habitat preference study. The two populations from northern France were collected by a colleague in the surroundings ofParis and Metz. Population refers to the populations used for electrophoresis. A - J: iC. alfacariensis, K-M: C. hyale. Population Site Location Lat Long Allozyme Field A del Commune 44 57.64 641.01 Poggio Italy X B+M Val de Pres French 44 58.47 6 30.33 Alps X X C+L Le Monetier les Bains French Alps 44 53.52 6 37.02 X X D Saint Pierre French Puy Alps 44 53.52 6 37.02 X E Saint French Zachery Alps 43 23.07 5 42.22 X F Ponteves Provence 43 33.55 6 02.53 X G Artigues Provence 43 36.05 5 47.54 X H Mi met Provence 43 24.54 5 30.19 X I Seine River outside Paris Provence X J+K of Surroundings Metz Northern France X Gero Northern France 44 59.03 6 28.56 X les Guibertes French 44 Alps 58.01 6 31.57 X le Cassset French 44 59.03 Alps 6 28.56 X Saint French 44 55.32 636.11 Chaffery Alps X le Rosier French Alps 44 56.02 6 40.58 X Villard Saint Pancrace French 44 52.09 Alps 6 38.05 X la Vachette French 44 Alps 54.58 6 39.55 X Laus de Cervieres French Alps 44 51,40 .6 43.35 X Granon French Alps 44 58.04 6 39.01 X Cervieres French 44 Alps 51.56 6 43.21 X for 6 minutes at 9000 x The trifuged g.
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