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© 1999 Macmillan Magazines Ltd letters to nature NATURE | VOL 401 | 9 SEPTEMBER 1999 | www.nature.com 157 was a high degree of band-sharing between individuals, probably due to past inbreeding in this population. The large number of monomorphic and common bands provided a reference ladder that allowed 28 apparently homologous polymorphic minisatellite fragments in the size range 4.0±23.0 kilobases (kb) to be identi®ed between ®ngerprints, thus enabling the ready comparison of all individuals. The order of the samples was randomized within and between gels, and the band patterns were scored by an assistant who had no knowledge of the lek sites of the individual birds. Eight free-ranging full-trained displaying lek males whose mating success varied were removed from Whipsnade Park during February 1991 and transferred to a peacock farm in Norfolk, UK. The peacocks were housed in separate pens and four naive adult peahens, known to be at least 2 years old, provided by the farm, were measured and randomly assigned to each pen on 14 March. Pens were checked daily for eggs (it was not possible to know which of the four hens laid which egg unless egg laying was observed) and any eggs found were labelled and removed. Groups of eggs originating from several different pens over several dates were mixed and placed under broody domestic chickens for incubation. Eggs were removed from the hens after 26 days and placed in a hatcher in batches, where each egg had its own compartment; each of the hatched chicks was given an individual colour ring combination. Each batch of eggs was incubated and hatched separately at approximately weekly intervals from May to August. Each batch of chicks was provided with a heat lamp and food and water ad libitum; batches were subsequently pooled and reared together. Females produced 519 eggs and the growth of the surviving 349 offspring was monitored. In January and February 1992, 12 offspring (7 males and 5 females) from each of the 8 males (3 from each of the 4 females per male) were introduced into Whipsnade Park. A matched sample of young was chosen from each pen so that there were no overall signi®cant differences in hatching dates or weights of the offspring between fathers (at day 84, F 7;95 0:838, P 0:559; at introduction, F 7;95 0:358, P 0:924). Care was taken to release the offspring in batches of eight, consisting of one young of the same sex from each pen. The fate of the offspring was recorded by a ®eld assistant who had no knowledge of the relatedness of any of the individuals, and the birds were observed every spring until they established permanent display sites in 1995 (aged 4). Of the introduced males, 19 were observed to have established permanent display sites in 1995 (Fig. 3). Mantel tests 7 that randomized the pairwise physical distances or band-sharing values, respectively, were performed on square-root transformed distances using the program RT v2.1 (ref. 21). We analysed nearest-neighbour associations using a program that rando- mized (100,000 times) the positions of relatives and non-relatives, as appropriate, and counted the number of occasions on which nearest neighbours were relatives. Received 11 March; accepted 2 July 1999. 1. Ho Èglund, J. & Alatalo, R. V. Leks (Princeton Univ. Press, Princeton, 1995). 2. Kokko, H., Lindstro Èm, J. Kin selection and the evolution of leks: whose success do young males maximise? Proc. R. Soc. Lond. B 263, 919±923 (1996). 3. Kokko, H., Sutherland, W. J., Lindstro Èm, J., Reynolds, J. D. & Mackenzie, A. Individual mating success, lek stability, and the neglected limitations of statistical power. Anim. Behav. 56, 755±762 (1998). 4. Alatalo, R. V., Ho Èglund, J., Lundberg, A. & Sutherland, W. J. Evolution of black grouse leksÐfemale preferences bene®t males in larger leks. Behav. Ecol. 3, 53±59 (1992). 5. Widemo, F. & Owens, I. P. F. Lek size, male mating skewand the evolution of lekking. Nature 373, 148±151 (1995). 6. Petrie, M., Halliday, T. & Sanders, C. Peahens prefer peacocks with elaborate trains. Anim. Behav. 41, 323±332 (1991). 7. Manly, B. F. Randomisation, Bootstrap and Monte Carlo Methods in Biology 2nd edn (Chapman & Hall, London, 1997). 8. Bruford, M. W., Hanotte, O., Brook®eld,J. F. Y. & Burke,T. in Molecular Genetic Analysis of Populations: A Practical Approach 2nd edn (ed. Hoelzel, A. R.) 287±336 (IRL, Oxford, 1998). 9. Petrie, M. Improved growth and survival of offspring of peacocks with more elaborate trains. Nature 371, 598±599 (1994). 10. Ho Èglund, J., Alatalo, R. V., Lundberg, A., Rintama Èki, P. T. & Lindell, J. Microsatellite markers reveal the potential for kin selection on black grouse leks. Proc. R. Soc. Lond. B 266, 813±816 (1999). 11. Lacy, R. C. & Sherman, P. W. Kin recognition by phenotype matching. Am. Nat. 121, 489±512 (1983). 12. Grosberg, R. K. & Quinn, J. F. The genetic control and consequences of kin recognition by the larvae of a colonial marine invertebrate. Nature 322, 457±459 (1986). 13. Potts, W. K., Manning, C. J. & Wakeland,E. K. Mating patterns in semi-natural populations of mice in¯uenced by MHC genotype. Nature 352, 619±621 (1991). 14. Getz, W. M. & Smith, K. B. Genetic kin recognition: honey bees discriminate between full and half sisters. Nature 302, 148 (1983). 15. Grafen, A. Do animals really recognise kin? Anim. Behav. 39, 42±54 (1990). 16. Alexander, R. D. Epigenetic rules and Darwinian algorithms: the adaptive study of learning and development. Ethol. Sociobiol. 11, 241±303 (1990). 17. Sherman, P. W. Multiple mating and kin recognition by self-inspection. Ethol. Sociobiol. 12, 377±386 (1991). 18. Sherman, P. W., Reeve, H. K. & Pfennig, D. W. in Behavioural Ecology: An Evolutionary Approach, 4th edn (eds Krebs, J. R. & Davies, N. B.) 69±97 (Blackwell Science, Oxford, 1997). 19. Blaustein, A. R., Bekoff, M. & Daniels, T. J. in Kin Recognition in Animals (eds Fletcher, D. J. C. & Michener, C. D.) 287±332 (Wiley, New York, 1987). 20. Hanotte, O., Burke, T., Armour, J. A. L. & Jeffreys, A. J. Hypervariable minisatellite DNA sequences in the Indian peafowl Pavo cristatus. Genomics 9, 587±597 (1991). 21. Manly, B. F. J. RT, A Program for Randomization Testing, version 2.1. (Centre for Applied Statistics and Mathematics, University of Otago, 1997). Acknowledgements This study was initiated while M.P. was at the Department of Zoology, University of Oxford and A.K. and T.B. were at the Department of Biology, University of Leicester. We thank the Zoological Society of London for permission to study the peafowl at Whipsnade Wild Animal Park; L. Lovett for help at Whipsnade; Q. Spratt for allowing M.P. To work at his peacock farm; A. Williams for help at the farm; P. Carpenter for help in scoring gels; E. Bell and A. Askew for writing computer programs; J. Brook®eld for advice; J. Futter for help with producing Figures; B. Hatchwell, M. Young, G. Roberts, F. Ratnieks and P. Watt for comments on the manuscript; and the NERC and the BBSRC for ®nancial support. Correspondence and requests for materials should be addressed to M.P. (e-mail: marion.petrie@ncl. ac.uk). ................................................................. An epigenetic mutation responsible for natural variation in ¯oral symmetry Pilar Cubas*, Coral Vincent & Enrico Coen John Innes Centre, Colney Lane, Norwich NR4 7UH, UK .............................................................................................................................................. Although there have been many molecular studies of morpho- logical mutants generated in the laboratory, it is unclear how these are related to mutants in natural populations, where the constraints of natural selection and breeding structure are quite different. Here we characterize a naturally occurring mutant of Linaria vulgaris, originally described more than 250 years ago by Linnaeus 1±3 , in which the fundamental symmetry of the ¯ower is changed from bilateral to radial. We show that the mutant carries a defect in Lcyc, a homologue of the cycloidea gene which controls dorsoventral asymmetry in Antirrhinum 4 . The Lcyc gene is exten- sively methylated and transcriptionally silent in the mutant. This modi®cation is heritable and co-segregates with the mutant phenotype. Occasionally the mutant reverts phenotypically during somatic development, correlating with demethylation of Lcyc and restoration of gene expression. It is surprising that the ®rst natural morphological mutant to be characterized should trace to methylation, given the rarity of this mutational mechan- ism in the laboratory. This indicates that epigenetic mutations may play a more signi®cant role in evolution than has hitherto been suspected. Mature wild-type ¯owers of Linaria vulgaris (toad¯ax) have ®ve petals that are united for part of their length to form a corolla tube ending in ®ve separate lobes (Fig. 1c, d). Dorsoventral asymmetry is clearly evident in the shape and colour of the petals. The two dorsal (adaxial) petals have relatively long strap-shaped lobes; the two lateral petals have wider lobes with a partially orange lip; and the ventral (abaxial) petal has a small lobe with an orange lip, and a spur-shaped nectary at its base. Dorsoventral asymmetry is also evident in the stamens: the dorsal stamen is arrested early in development to give a sterile staminode (Fig. 1d), and the two lateral stamens are shorter and less hairy than the two ventral stamens. Flowers of naturally occurring peloric mutants in Linaria are radially symmetrical (Fig. 1a±d). All ®ve petals resemble the ventral petal of wild type, each having a small lobe with an orange lip, and a spur at their base. Similarly, there are ®ve stamens, all of which closely resemble the ventral stamens of wild type in length and hairiness. In being fully ventralized, these mutant Linaria ¯owers resemble peloric mutants of Antirrhinum which lack the activity of two related genes, cycloidea (cyc) and dichotoma 4,5 . The peloric mutation in Linaria is recessive, as crosses to wild type yielded essentially wild-type F 1 progeny. Only one of the F 1 individuals occasionally gave one or two extra spurs. We compared the development of wild-type and peloric ¯owers of Linaria by scanning electron microscopy. No differences were *Present address: Centro Nacional de Biotecnologia Campus de la Universidad Auto  noma de Madrid, Cantoblanco, 28049, Madrid, Spain.
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