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A.J. Bateman (1948) hypothesized that a metric of sexual selection is in sex differences of intrasexual variance in number of mates (V NM). AJB predicted that (a) males have greater variance in reproductive success (V RS) than females; (b) males have greater V NM than females; and (c) a positive relationship between V NM and V RS is stronger among males. AJB used phenotypically observable mutations in offspring to identify parents and to count subjects' NM and RS. AJB's conclusions matched his predictions, later called "Bateman's Principles." Empirical challenges to his conclusions guided analyses herein. (a) AJB's analysis pseudo-replicated sample sizes, violating a sexual selection assumption: That is, individuals must be in the same population to choose and compete. (b) AJB's methods overestimated subjects with no mates while underestimating subjects with one or more. (c) A replication (Gowaty et al., 2012) showed that offspring inheriting nametags from both parents often died before expressing adult phenotypes, proving some of AJB's methods produced biased data. Science historian Thierry Hoquet located AJB's archived, handwritten laboratory notes, photocopied, and transcribed them. We tested each of the 65 unique populations for expected combinations in offspring of parental mutations: 41.5% failed Punnett's tests: Offspring carrying nametags simultaneously from both parents were missing showing estimates of parents' NM and V NM were undercounted. 58.5% of populations met Punnett's expectations providing an unparalleled opportunity to re-evaluate AJB's predictions. 34 unbiased populations had no sex differences in V RS; 37 had no sex differences in V NM. No sex differences in slopes of RS and NM occurred in any unbiased population. Regressions showed weak, positive, significant associations between V NM and V RS for females and males, contrary to AJB's prediction that the relationship would be positive in males but not in females. AJB's laboratory data are inconsistent with "Bateman's Principles."

Bateman's experimental study of Drosophila melanogaster produced conclusions that are now part of the bedrock premises of modern sexual selection. Today it is the most cited experimental study in sexual selection, and famous as the first experimental demonstration of sex differences in the relationship between number of mates and relative reproductive success. We repeated the experimental methodology of the original to evaluate its reliability. The results indicate that Bateman's methodology of visible mutations to assign parentage and reproductive success to subject adults is significantly biased. When combined in offspring, the mutations decrease offspring survival, so that counts of mate number and reproductive success are mismeasured. Bateman's method overestimates the number of subjects with no mates and underestimates the number with one or more mates for both sexes. Here we discuss why Bateman's paper is important and present additional analyses of data from our monogamy trials. Monogamy trials can inform inferences about the force of sexual selection in populations because in monogamy trials male-male competition and female choice are absent. Monogamy trials also would have provided Bateman with an a priori test of the fit of his data to Mendel's laws, an unstated, but vital assumption of his methodology for assigning parentage from which he inferred the number of mates per individual subject and their reproductive success. Even under enforced monogamous mating, offspring frequencies of double mutant, single mutant and no mutant offspring were significantly different from Mendelian expectations proving that Bateman's method was inappropriate for answering the questions he posed. Double mutant offspring (those with a mutation from each parent) suffered significant inviability as did single mutant offspring whenever they inherited their mother's marker but the wild-type allele at their father's marker locus. These inviability effects produced two important inaccuracies in Bateman's results and conclusions. (1) Some matings that actually occurred were invisible and (2) reproductive success of some mothers was under-estimated. Both observations show that Bateman's conclusions about sex differences in number of mates and reproductive success were unwarranted, based on biased observations. We speculate about why Bateman's classic study remained without replication for so long, and we discuss why repetition almost 60 years after the original is still timely, necessary and critical to the scientific enterprise. We highlight overlooked alternative hypotheses to urge that modern tests of Bateman's conclusions go beyond confirmatory studies to test alternative hypotheses to explain the relationship between mate number and reproductive success.

Polyandry is a paradox: why do females mate multiple times when a single ejaculate often provides enough sperm for lifetime egg production? Gowaty et al. addressed explanations for polyandry in Drosophila pseudoobscura from the perspective of hypotheses based on sex differences in costs of reproduction (CoR). Contrary to CoR, Gowaty et al. showed that (1) a single ejaculate was inadequate for lifetime egg production; (2) polyandry provided fitness benefits to females beyond provision of adequate sperm and (3) fitness benefits of polyandry were not offset by costs. Here, I discuss predictions of the ad hoc hypotheses of CoR and three alternative hypotheses to CoR to facilitate a discussion and further development of a strong inference approach to experiments on the adaptive significance of polyandry for females. Each of the hypotheses makes testable predictions; simultaneous tests of the predictions will provide a strong inference approach to understanding the adaptive significance of multiple mating. I describe a sex-symmetric experiment meant to evaluate variation in fitness among lifelong virgins (V); monogamous females and males with one copulation (MOC); monogamous females and males with multiple copulations (MMC); PAND, polyandrous females; and PGYN, polygynous males. Last, I recommend the study of many different species, while taking care in choice of study species and attention to the assumptions of specific hypotheses. I particularly urge the study of many more Drosophila species both in laboratory and the wild to understand the "nature of flies in nature," where opportunities and constraints mold evolutionary responses.

Western burrowing owls (Athene cunicularia hypugaea) are ground-dwelling owls distributed throughout western North America. Because of population declines, this species is considered endangered in Canada, and burrowing owls are listed as a species of conservation concern in states of the western USA. Korfanta et al. (2002) previously presented primers for seven microsatellite loci in burrowing owls. Parentage and relatedness studies require a larger number of markers for accuracy and precision. Here, we developed and characterized 18 additional microsatellite DNA loci, and we tested these loci in 23 individuals. The number of alleles per locus ranged from 2 to 11; two loci deviated from Hardy–Weinberg equilibrium following Bonferroni correction; we did not detect linkage disequilibrium following Bonferroni correction; and the probability of exclusion for parent pairs using all loci was >0.9999. We envision these loci will facilitate detailed analyses of the genetic mating system of burrowing owls, which is poorly understood.

Blue-footed boobies (Sula nebouxii) are socially monogamous, colonial seabirds exhibiting intra-specific nest parasitism and extra-pair copulations. Prior DNA fingerprinting assays failed to detect extra-pair offspring in the nests of congeners, and the rate of intra-specific nest parasitism has not been estimated using molecular techniques. We describe the development and characterization of 11 microsatellite DNA loci, tested using 31 individuals collected on Isla Isabel, Nayarit, México. The number of alleles per locus ranged from three to 22, averaging seven; total exclusionary power of the microsatellite panel was 0.99; no loci deviated from Hardy-Weinberg equilibrium; and we did not detect linkage disequilibrium following Bonferroni correction. This microsatellite panel will facilitate future studies of nest parasitism and extra-pair paternity in blue-footed boobies.