1. John Pannell Department of Plant Sciences University of Oxford Possible links between sexual-system evolution and demographic processes in plants and animals

3. Hermaphroditism Gynodioecy Androdioecy (Male-sterility) (Female-sterility)

4. Relative production of seeds or pollen (relative to hermaphrodites) Frequency of males or females Maintaining males versus females

5. Relative production of seeds or pollen (relative to hermaphrodites) Frequency of males or females Maintaining males versus females Gynodioecy under selfing & inbreeding depression

6. Relative production of seeds or pollen (relative to hermaphrodites) Frequency of males or females Maintaining males versus females Gynodioecy under selfing & inbreeding depression Androdioecy under selfing & inbreeding depression

7. Predictions 1.Androdioecy should be difficult to evolve 2.Androdioecy should occur only where hermaphrodites are prevented from selfing In fact… 1.Androdioecy has evolved several times 2.Males typically occur with partially selfing hermaphrodites

8. Mercurialis annua Datisca glomerata Schizopepon bryoniaefolius Occurrence of androdioecy Kryptolebias marmoratus

9. Several species of branchiopod crustaceans Sassaman, 1995 Herm Male Steve Weeks

10. Several species of the Oleaceae Wallander, 2001

11. Durand (1963) Mercurialis annua

12. Established population males hermaphrodites Immigration males selected hermaphrodites selected with female-biased sex allocation Colonisation population growth reduced selfing

13. Obbard, Harris & Pannell (Am Nat, 2006) Within-population diversity Pair-wise differentiation

14. 445 populations 5 transitions in breeding system 3 years of demographic sampling Males present Males absent

15. Males + females Hermaphrodites Dorken & Pannell (2007: Heredity)

16. Occupancy (% occupied sites) 405060708090100 Abundance (number of plants) 0 1000 2000 3000 4000 5000 6000 7000 Males absent Males present Eppley & Pannell (2006: American Naturalist)

17. Dorken, Freckleton & Pannell (unpublished data) 25/171 49/185 Males present Males absent

18. 2x Dioecy (West) Dioecy (Central) Dioecy (East) Monoecy Obbard, Harris & Pannell (American Naturalist, 2006) 6x

19. Dioecy (West) Dioecy (Central) Dioecy (East) Monoecy 6x Is there less inbreeding depression in northern populations of M. annua?

20. Pujol et al. (PNAS, 2009)

21. Dioecy (West) Dioecy (Central) Dioecy (East) Monoecy 6x Is there less quantitative genetic variation for sex allocation in northern populations of M. annua?

22. Pujol and Pannell (2008, Science)

23. hermaphrodites maleshermaphrodites Sex allocation Frequency 0.01.0

24. Male frequency Nutrient status

25. Dorken and Pannell (Current Biology, in press)

26. Durand (1963)

27. Korbecka and Pannell, unpubl. Selfing rates in different patches With males Without males

28. Dorken, Freckleton & Pannell (unpublished data) Eppley and Pannell (2008: Evolution)

29. Eulimnadia texana Datisca glomerata Schizopepon bryoniaefolius Occurrence of males with hermaphrodites Kryptolebias marmoratus Herm Male Caenorhabditis elegans

30. Weeks et al. (2006) Males + hermaphrodites 24–180 million years ago Males + females Hermaphrodites Males and hermaphrodites in Eulimnadia species Herm Male

31. Why is androdioecy in Eulimnadia so ancient? Hermaphrodites are the heterogametic sex W/Z Z/Z

32. W/Z Z/Z Deleterious recessives on W NOT expressed FIXED by drift W/Z Z/Z W/Z W/W Selfing produces homozygous W Load on W now expressed Fitness of W/W < W/Z Selection for reproductive assurance Females produce an ovotestis Androdioecy evolves

33. Male frequency Probability of finding a mate  = 0.9 = 0.9 = 0.1 (recessive load on W)  = 0.1 l = 0 (no recessive load on W)  = 0.9  = 0.5 l > 0 (recessive load on W) Maintenance of males

34. Male frequency Probability of finding a mate  = 0.9 = 0.9 = 0.1  = 0.1 l = 0 (no recessive load on W)  = 0.9  = 0.5 l > 0 (recessive load on W) Maintenance of males Pannell (2008: Genetical Research)

35. Males maintained by overdominance? ZZ males: low fitness –can’t find a mate WW hermaphrodites: low fitness –reproductive assurance –BUT recessive genetic load on W chromosome WZ hermaphrodites: high fitness –reproductive assurance –AND sheltering of genetic load on W chromosome

36. Verdu, Montilla & Pannell (Proc. Royal Soc., B, 2004) Fraxinus ornus Oleaceae family Dioecy & androdioecy are frequent in genus and family

37. First puzzle… Males and hermaphrodites co-occur Fraxinus ornus Implies androdioecy 1:1 sex ratio Implies cryptic dioecy Hermaphrodites do sire seeds Implies androdioecy Functional Ecology (2002): 16: 858-869 Proc. Royal Soc., B (2004): 271: 2017-2023 Evolution (2006)

38. father Male-sired seedlings grow 8% faster than hermaphrodite-sired seedlings Proc. Royal Soc., B (2004): 271: 2017-2023 mother

39. Fraxinus ornus Hermaphrodites can be fathers but not grandfathers Hermaphrodites are functionally female Proc. Royal Soc., B (2004): 271: 2017-2023 Evolution (2006)

40. Second puzzle… Females produce lots of pollen… Why? Fraxinus ornus F F M M Intense competition in the seed shadow

41. Second puzzle… Females produce lots of pollen… Why? Fraxinus ornus F F M M Intense competition in the seed shadow Rival’s seedlings are less competitive ESS: all females invest up to 50% of reproductive resources in pollen Pannell (unpubl.)

42. Third puzzle… What is the mechanism of sabotage?

43. Thanks to… Darren Obbard Richard Buggs Stephen Harris Sarah Eppley Marcel Dorken Paul Rymer Rob Freckleton Grazyna Korbecka Stephen Weeks … and many undergraduate assistants NERC Royal Society BBSRC European Union