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Chapter 13: Wholly Virgin
For general biology of bdelloid rotifers, see Donner (1966) and Ricci (1987). David Mark Welch kindly told me that the age of the bdelloid rotifers, based on
molecular evidence, is eighty-five million years. The problem of the evolution of sex has been discussed many times: see, for example, Maynard Smith (1978), Bell (1982), and Kondrashov (1993). For the particular problem posed by the bdelloid rotifers, see Maynard Smith (1986) and Judson and Normark (1996). Maynard Smith (1986) dubbed the bdelloid rotifers an “evolutionary scandal.” My description of the cost of sex is drawn from Maynard Smith (1978), page 3. My account of bacterial sex is based on Levin (1988), Maynard Smith et al. (1993), and Davies (1994); for viral sex, see Chao (1992). An account of meiosis can be found in any basic genetics textbook.
For other putative ancient asexuals, see Judson and Normark (1996), although for the age of the darwinulid ostracods, see Butlin et al. (1999). For the discrediting of the chaetonotid gastrotrichs, see Weiss and Levy (1979); of the aphids, see Normark (1999). Ben Normark kindly told me of the discovery of males. Hurst et al. (1992) and Little and Hebert (1996) cast doubt on the existence of ancient asexuals. For the pattern of molecular evolution that you expect to observe in an ancient asexual lineage, see Judson and Normark (1996) and Birky (1996). For proof that the bdelloid rotifers have the expected pattern and have therefore been without sex for millions of years, see Mark Welch and Meselson (2000).
The notion that asexuals cannot evolve is extremely common. For the number and general distribution of bdelloid species, see Ricci (1987); for evidence that bdelloids occur in the Antarctic, see Everitt (1981); for the lives of seisonid rotifers, see Ricci (1992). In arguing that most mutations are neutral, Miss
Philodina
is adopting the position of Kimura (1983).
Muller (1964) proposed the ratchet. It has been commented on extensively; for how asexuals can evade the ratchet, see Judson and Normark (1996). The hatchet mechanism was proposed by Kondrashov (1984) and Kondrashov (1988). For a general account of
Atta colombica
and the fungus, see Wilson (1971), pages 41—48; for evidence that the fungus is an ancient asexual, see Chapela et al. (1994), especially note 19; for all details of
Escovopsis,
see Currie et al. (1999). The idea that parasites are important in maintaining sex and variation was suggested by Haldane (1949), Bremermann (1980), Hamilton (1980), and Tooby (1982); Bell (1982) christened this theory the Red Queen. See Carroll (1871) for the Red Queen quotation. For clonal reproduction in armadillos, see Loughry et al. (1998); for the armadillo's penis, see Wassersug (1997). Ladle et al. (1993) suggested ancient asexuals can escape the Red Queen if they disperse without parasites. For the survival of bdelloid rotifers after anhydrobiosis, see Ricci (1987).
 
Postscript
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