So Bummed Out in Berkeley
For self-restraint among male California mice, see Gubernick and Nordby (1993); for monogamy in California mice, see Ribble (1991). For excellent overviews of the hormonal mechanisms of monogamy, see Young et al. (1998) and Insel and Young (2001). For the role of vasopressin in prairie vole monogamy, see Winslow et al. (1993); for transgenic mice and vasopressin, see Young et al. (1999); for behavioral contrasts with montane voles, see Shapiro and Dewsbury (1990); for hormonal contrasts, see Young et al. (1998). For frequent sex in the Indian crested porcupine, see Sever and Mendelssohn (1988); in the wood roach, see Nalepa (1988). For nonidentical twins having different fathers, see Phelan et al. (1982). For size differences between male and female southern elephant seals, see Nowak (1999), page 880; between male and female gorillas, see Nowak (1999), page 620. For differences in testicle size between humans and other apes, see Short (1979). For an excellent discussion of the fiction of high rates of infidelity as measured by paternity testing in humans, see Macintyre and Sooman (1991); for studies showing low rates of infidelity, see Ashton (1980) and Sasse et al. (1994); for the rate of 11.8 percent, see Cerda-Flores et al. (1999); for infidelity among Sykeses, see Sykes and Irven (2000). For a genetic predisposition toward promiscuity in crickets, see Solymar and Cade (1990); in fruit flies, see Pyle and Gromko (1981).
Chapter 11: The Fornications of Kings
Lesser mealworm beetle Alphitobius diaperinus
White-lipped land snail Triodopsis albolabris
True armyworm moth Pseudaletia unipuncta
Woodlouse Armadillidium vulgare
Wood lemming Myopus schisticolor
House sparrow Passer domesticus
Great land crab Cardisoma guanhumi
Pink salmon Oncorhynchus gorbuscha
Soapberry bug Jadera haematoloma
Soapberry tree Sapindus saponaria
Round-podded golden rain tree Koelreuteria paniculata
Aghast in Arkansas
For the natural history of Acarophenax mahunkai, see Steinkraus and Cross (1993). Many thanks to Don Steinkraus for confirming that the mites’ ability to detect the sex of beetles is probable but unknown. For the effects of close incest in humans, see Seemanová (1971). For a general account of inbreeding depression and recessive genes, see any population genetics textbook; for a more technical account of the subject (including discussions of alternative, but less widely accepted explanations for inbreeding depression that are not mentioned by Dr. Tatiana), see Charlesworth and Charlesworth (1987). For marriages on kibbutzim, see Shepher (1971). For self-copulation in Dendrobaena rubida, see Andre and Davant (1972). For the evolution of selfing rates in hermaphrodites, see Jarne and Charlesworth (1993); for emergency selfing in the white-lipped land snail, see McCracken and Brussard (1980). For Hawaiian incest, see Malo (1903), chapter 18; for Egyptian incest, see Tyldesley (1994), pages 198–99; for a general discussion of royal incest (including the notion that it is a result of stratification) and for incest among the Incas, see van den Berghe and Mesher (1980). For the independent origins of haplodiploidy and paternal genome elimination, see Mable and Otto (1998). For haplodiploidy and inbreeding in pinworms, see Adamson (1989); in insects and mites (and for discussions of paternal genome elimination), see Wrensch and Ebbert (1993). For incest in the button beetle, see Hamilton (1993), page 430. For the life and times of Scleroderma immigrans, see Wheeler (1928), pages 62–64. The advantage of having haploid males when making a switch to incest has been remarked on by many authors; for a formal treatment, see Werren (1993).
Piqued in Darien
For a marvelous account of the biology of the moth ear mite, see Treat (1975), chapter 7; for the sex ratio in moth ear mites, as well as proof of their paternal genome elimination, see Treat (1965). The moth nursery rhyme is original to Dr. Tatiana. For mites on the antennae and on the feet of army ants, for hummingbird nostril mites, and for fruit bat eyeball mites, see Walter and Proctor (1999), pages 200–201 (Dave Walter tells me that some species of army ant are likely to have both types of mite); for human follicle mites, see Nutting (1976); for quill mites, see Kethley (1971). For Fisher’s argument on sex ratios, see Fisher (1999), pages 141–43. For the effect of war on the human sex ratio, see Graffelman and Hoekstra (2000). For feminization of genetic males in woodlice, see Bull (1983), page 200; for sex ratio skew in wood lemmings, see Nowak (1999), pages 1482–83; for their maverick chromosomes, see Bull (1983), pages 79-80. For sex ratios in quill mites, see Kethley (1971). For sex ratios in Acarophenax mahunkai, see Steinkraus and Cross (1993). For the evolution of sex ratios under inbreeding, see Hamilton (1996), chapter 4. For selfing rates and allocation to male tissue in Utterbackia imbecillis, see Johnston et al. (1998); for smaller flowers in selfing plants, see Charnov (1982), pages 261–68; for runty males in incestuous animal species, see Hamilton (1993). For sex ratio adjustment in Nasonia vitripennis, see Werren (1980). For control of sex ratio under paternal genome elimination in Typhlodromus occidentalis, see Nagelkerke and Sabelis (1998).
Gagging for It in Florida
For mangrove fish sharing burrows with land crabs and for their ability to survive out of water, see Taylor (1990). For outbreeding depression in pink salmon, see Gharrett et al. (1999). My discussion of the soapberry bug is a simplification of Carroll and Boyd (1992). For male frequencies in Rivulus marmoratus, see Taylor et al. (2001); for effects of outcrossing, see Taylor (2001).
Chapter 12: Eve’s Testicle
Black hamlet fish Hypoplectrus nigricans
Green spoon worm Bonellia viridis
Spotted hyena Crocuta crocuta
Brown hyena Hyaena brunnea
Striped hyena Hyaena hyaena
Aardwolf Proteles cristatus
Wildebeest Connochaetes taurinus
Thomson’s gazelle Gazella thomsonii
Paper nautilus Argonauta argo
Green poison arrow frog Dendrobates auratus
Spraying characid Copeina arnoldi
Dayak fruit bat Dyacopterus spadiceus
Looking for a Baker’s Dozen in the Forests of Romania
For an excellent description of the life cycle of Physarum polycephalum, see Bailey (1997); for the system of sexes, see Kawano et al. (1987) and Bailey (1997); for matA and the removal of the mitochondria after fusion, see Meland et al. (1991). My definition of sexes follows standard practice. For predictions that if finding a mate is difficult, isogamous organisms should have a large number of sexes, see Iwasa and Sasaki (1987); for the notion that having two sexes is deeply strange, see Hurst (1996); for the efficiency of inbreeding prevention and outbreeding promotion in organisms with multiple mating types, see Raper (1966). For problems evolving from zero to two sexes, see Hoekstra (1987) and Hutson and Law (1993). For the notion that sexes evolved to prevent conflict between cytoplasmic elements, see Eberhard (1980), Cosmides and Tooby (1981), Hurst and Hamilton (1992), Hurst (1996). For mitochondria and chloroplast transmission in Chlamydomonas reinhardtii, see Gillham (1994). For large numbers of sexes in mushrooms and ciliates, see references in Hurst and Hamilton (1992); for twenty thousand sexes in Schizophyllum commune, see Kothe (1996); for difficulties in regulating the transmission of genetic elements when there are more than two sexes, see Hurst (1996); for badly behaved slime mold mitochondria, see Kawano et al. (1991).
Ready to Litigate in Tallahassee
The problem of evolving males and females from isogamy has been discussed extensively For a recent review, see Randerson and Hurst (2001). For Chlamydomonas moewusii producing small cells, see Weise et al. (1979). For the advantage of being a male, see Parker et al. (1972)—for problems with the theory, see Randerson and Hurst (2001). In arguing that bigness is an advantage because it makes you easier to find, I follow Dusenbery (2000). For the notion that males and females evolved to control cytoplasmic elements, see Hastings (1992); for problems with this idea, see Randerson and Hurst (2001).
Group Sexists in Santa Catalina
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p; For orgies in Aplysia californica, see Pennings (1991); for spawning behavior of the black hamlet fish, see Fischer (1980). For fusion in Diplozoon gracile, see Justine et al. (1985). For aerial sex in Limax maximus, see Langlois (1965); in Limax redii, see Baur (1998). For a superb general treatment of when to be a hermaphrodite, see Charnov (1982); for low density and hermaphrodites, see Ghiselin (1969); for the prediction that wind pollination leads to separate sexes, see Charnov et al. (1976). For wind-pollinated plants tending to having separate sexes and pollinator-pollinated plants tending toward hermaphroditism, see Renner and Ricklefs (1995). For hermaphroditism in fish, see Charnov (1982), table 12.1; for deep-sea comb jellies with separate sexes, see Harbison and Miller (1986); for patterns of hermaphroditism in bivalves and barnacles, see Charnov (1982), page 239; for the absence of hermaphrodites from many groups, see Ghiselin (1974), chapter 4. Little has been written about the transition between separate sexes and hermaphroditism. For some of the genetic considerations, see White (1973). For males, females, and hermaphrodites in the cactus Pachycereus pringlei, see Fleming et al. (1994).
Too Much Heavy Breathing near Malta
For dwarf males and sex determination in the green spoon worm, see Jaccarini et al. (1983). Many thanks to René Hessling for calculating the size difference between males and females. For a general overview of sex determination, see Bull (1983); for sex chromosomes, see particularly pages 16–20; for patterns of sex determination in reptiles, see pages 115–22. For multiple independent origins of males hatching from unfertilized eggs, see Mable and Otto (1998), table 1; for sex determination by fungal infection in Stictococcus sjoestedti, see Bacci (1965), pages 154–55. For Capitella becoming a hermaphrodite at low density, see Holbrook and Grassle (1984). For sex change in the slipper limpet, see Hoagland (1978); in Ophryotrocha puerilis, see Berglund (1986). For a general account of when to change sex, see Charnov (1982); for the size advantage model of sex change, see Ghiselin (1969). For examples of dwarf males in other species and for the notion that dwarf males are favored when females are both sedentary and few and far between, see Ghiselin (1974), chapter 7; for dwarf males in anglerfish, and for general anglerfish biology see Bertelsen (1951); for the claim that anglerfish males have the largest noses in proportion to their bodies, see Andersson (1994), page 257.
Don’t Wanna Be Butch in Botswana
For the hunting and dining habits of spotted hyenas and for lions scavenging from hyenas, see Kruuk (1972), especially chapter 5; for comparisons with other members of the hyena family, see Estes (1991), chapter 20, and Nowak (1999), pages 786–93. For the social structure of spotted hyenas, see Kruuk (1972), chapter 6, and Frank (1997). For the belief that the spotted hyena was a hermaphrodite and for the structure of the female’s genitalia, see Kruuk (1972), page 210; for copulation in the spotted hyena, see Frank (1997); for the structure of the birth canal and for estimated death rates during birth, see Frank et al. (1995). For discussion of the idea that the female’s phallus has evolved because of its use in greeting ceremonies, see Kruuk (1972), chapter 6, and Frank (1997); for the theory that the phallus is an antirape device, see East et al. (1993); for a vigorous debunking of the antirape theory, see Frank (1997). For aggressiveness in female mice and its relationship to position in the womb, see vom Saal (1989); for androgen exposure and genital abnormalities, see Frank (1997). For siblicide in spotted hyenas, see Frank et al. (1991); for greater reproductive success of dominant females, see Frank (1997). For the blocking of androgen exposure in the womb and its lack of effect on female genitalia in hyenas, see Drea et al. (1998). For spiders charging their pedipalps, see Bristowe (1958), pages 65-67; for genitalia in the seahorse, see Eberhard (1985), page 68. For the genitalia of Sapha amicorum, see Marcus (1959). For the penis of the paper nautilus, see Müller (1853) and Young (1959). For the pouch in the leech Marsupiobdella africana, see van der Lande and Tinsley (1976); for hunting in Helobdella striata, see Kutschera and Wirtz (1986). For parental care in the green poison arrow frog, see Wells (1978) and Summers (1989). For the spraying characid laying and tending eggs, see Krekorian (1976). For male Dayak fruit bats producing milk, see Francis et al. (1994). The gender bender poem is original to Dr. Tatiana.
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
For the idea that sex arose through cannibalism, see Margulis and Sagan (1986); for the notion that it arose through the action of genetic elements trying to promote their own spread, see Hickey and Rose (1988); for the notion that it arose to enable DNA repair, see Bernstein et al. (1988).
BIBLIOGRAPHY
Abele, L. G., and S. Gilchrist, 1977. Homosexual rape and sexual selection in Acanthocephalan worms. Science 197: 81–83.
Adamson, M. L., 1989. Evolutionary biology of the oxyurida (Nematoda): Bio-facies of a haplodiploid taxon. Advances in Parasitology 28: 175–228.
Afzelius, B. A., B. Baccetti, and R. Dallai, 1976. The giant spermatozoon of Notonecta. Journal of Submicroscopic Cytology 8: 149–61.
Afzelius, B. A., and R. Dallai, 1983. The paired spermatozoa of the marine snail, Turritella communis Lamarck (Mollusca, Mesogastropoda). Journal of Ultrastructure Research 85: 311–19.
Aguadé, M., 1999. Positive selection drives the evolution of the Acp29AB accessory gland protein in Drosophila. Genetics 152: 548–51.
Alatalo, R. V, J. Höglund, A. Lundberg, and W J. Sutherland, 1992. Evolution of black grouse leks: Female preferences benefit males in larger leks. Behavioral Ecology 3: 53–59.
Alcock, J., 1996. The relation between male body size, fighting, and mating success in Dawson’s burrowing bee, Amegilla dawsoni (Apidae, Apinae, Antho-phorini). Journal of Zoology 239: 663–74.
Alcock, J., G. C. Eickwort, and K. R. Eickwort, 1977. The reproductive behavior of Anthidium maculosum (Hymenoptera: Megachilidae) and the evolutionary significance of multiple copulations by females. Behavioral Ecology and Sociobiology 2: 385–96.
Aldrich, J. M., and L. A. Turley, 1899. A balloon-making fly. American Naturalist 33: 809–12.
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