In a variety of fish species with paternal care of offspring, females prefer to spawn in nests that already contain eggs. This female preference has been hypothesized to explain egg thievery in male sticklebacks, allopaternal care of eggs in minnows, and the evolution of egg-mimicking body features in male cichlids and darters. Here we employ microsatellite-based parentage analyses to evaluate the reproductive success of striped darter (Etheostoma virgatum) males that appear to utilize two of these functionally related tactics to entice females to spawn in their nests. In an isolated population (Clear Creek, Ky.), we observed that breeding males develop conspicuous white spots on their pectoral fins. If these spots are egg mimics, as we suspect, then this represents the fourth independent evolutionary origin of egg mimicry documented to date in darters, the first based on pigmentation (as opposed to physical structures), and the first in which the egg mimics vary greatly in number among males. From direct counts of microsatellite genotypes in clutches of embryos, at least 3.8 females contributed to the progeny within a typical nest, and females tended to spawn preferentially with males that were larger and displayed more egg-mimic spots. In another population (Hurricane Creek, Tenn.) without egg mimics, the multi-locus genetic data document that allopaternal care is common, especially among the smallest males who sometimes tend nests containing their own as well as an earlier sire's offspring. Thus, these foster males had adopted egg-containing nests and then successfully spawned with subsequent females. Overall, the genetic data on paternity and maternity, in conjunction with field observations, suggest that egg mimicry and allopaternal care are two mate-attracting reproductive tactics employed by striped darter males to exploit female preferences for spawning in nests with 'eggs'.

We address various statistical aspects of biological parentage in multi-offspring broods that arise via multiple paternity or multiple maternity and, hence, consist of mixtures of full- and half-sibs. Conditioned on population genetic parameters, computer simulations described herein permit estimation of: (1) the mean number of offspring needed to detect all parental gametes in a brood and (2) the relationship between the number of distinct parental gametes found in a brood and the number of parents. Results are relevant to the design of empirical studies employing molecular markers to assess genetic parentage in polygynous or polyandrous species with large broods, such as are found in many fishes, amphibians, insects, plants and other groups. The utility of this approach is illustrated using two empirical data sets.

Understanding how variation in reproductive success is related to demography is a critical component in understanding the life history of an organism. Parentage analysis using molecular markers can be used to estimate the reproductive success of different groups of individuals in natural populations. Previous models have been developed for cases where offspring are random samples from the population but these models do not account for the presence of full- and half-sibs commonly found in large clutches of many organisms. Here we develop a model for comparing reproductive success among different groups of individuals that explicitly incorporates within-nest relatedness. Inference for the parameters of the model is done in a Bayesian framework, where we sample from the joint posterior of parental assignments and fertility parameters. We use computer simulations to determine how well our model recovers known parameters and investigate how various data collection scenarios (varying the number of nests or the number of offspring) affect the estimates. We then apply our model to compare reproductive success among different age groups of mottled sculpin, Cottus bairdi, from a natural population. We demonstrate that older adults are more likely to contribute to a nest and that females in the older age groups contribute more eggs to a nest than younger individuals.

Statistical methods for estimating genetic parentage are increasingly applied to accommodate limited marker polymorphism and the incomplete sampling of individuals. Neff et al. (2000a, Mol. Ecol. 9, 515-528; 2000b, Mol. Ecol. 9, 529-539) published a method (Pat) that estimates the proportion of next-generation individuals sired by a focal male, taking into account that the male may be genetically compatible, by random chance, with offspring that are not his own. Here we employ this method to reestimate paternity of 68 nest-guarding males from several fish species. The difference between the conventional exclusion-based estimate and Pat was >0.05 in only four of the 68 (5.9%) fish nests analyzed. An analytical formula shows that the difference between the two estimates is expected to be negligible if the focal male is consistent with a large proportion of the genotyped offspring, or if marker polymorphism is high. In addition, computer simulations illustrate how numbers of marker loci and their levels of genetic polymorphism, as well as the mating system of the organism under study, can influence estimates of paternity derived from exclusion-based estimates and Pat. Finally, we discuss various applications of these estimators including cases where additional biological information is present in the form of behavioral observations on parental care.

Most genetic surveys of parentage in nature sample only a small fraction of the breeding population. Here we apply microsatellite markers to deduce the genetic mating system and assess the reproductive success of females and males in an extensively collected, semi-closed stream population of the mottled sculpin fish, Cottus bairdi. In this species, males guard nest rocks where females deposit the eggs for fertilization. The potential exists for both males and females to mate with multiple partners and for males to provide parental care to genetically unrelated offspring. Four hundred and fifty-five adults and subadults, as well as 1,259 offspring from 23 nests, were genotyped at five polymorphic microsatellite loci. Multilocus maternal genotypes, deduced via genetic analyses of embryos, were reconstructed for more than 90% of the analysed nests, thus allowing both male and female reproductive success to be estimated accurately. There was no genetic evidence for cuckoldry, but one nest probably represents a takeover event. Successful males spawned with a mean of 2.8 partners, whereas each female apparently deposited her entire clutch of eggs in a single nest (mean fecundity = 66 eggs/female). On average, genetically deduced sires and dams were captured 1.6 and 9.3 metres from their respective nests, indicating little movement by breeders during the spawning season. Based on a 'genetic mark-recapture' estimate, the total number of potentially breeding adults (c. 570) was an order-of-magnitude larger than genetically based estimates of the effective number of breeders (c. 54). In addition, significantly fewer eggs per female were deposited in single than in multidam nests. Not only were perceived high-quality males spawning with multiple partners, but they were receiving more eggs from each female.

Supplemental breeding is an intensive population management strategy wherein adults are captured from nature and spawned in controlled settings, and the resulting offspring are later released into the wild. To be effective, supplemental breeding programs require crossing strategies that maximize offspring production while maintaining genetic diversity within each supplemental year class. We used computer simulations to assess the efficacy of different mating designs to jointly maximize offspring production and maintain high levels of genetic diversity (as measured by the effective population size) under a variety of biological conditions particularly relevant to species with high fecundity and external fertilization, such as many fishes. We investigated four basic supplemental breeding designs involving either monogamous pairings or complete factorial designs (in which every female is mated to every male and vice versa), each with or without the added stipulation that all breeders contribute equally to the tolal reproductive output. In general, complete factorial designs that did not equalize parental contributions came closest to the goal of maximizing offspring production while still maintaining relatively large effective population sizes. Next, we estimated the effective population size of 10 different supplemental year classes within the breeding program of the robust redhorse (Moxostoma robustum). Two year classes failed to produce progeny, whereas successful year classes used partial factorial designs to realize effective sizes ranging from 2 to 26 individuals. On average, a complete factorial design could increase the effective size of each robust redhorse supplemental year class by 19%.