Richard D. Howard

Individual variation in male vocal traits and female mating preferences inBufo americanus

We investigated the amount of variation in mating behaviour between and within individual male and female American toads, because both sources of trait variation can influence the course of sexual selection. Males varied in all four call parameters investigated (dominant call frequency, pulse rate, call rate and call duration). Individual males lowered the dominant frequency of their call when they interacted vocally with nearby males. Dominant call frequency was more highly correlated with body size in vocally interacting males than in non-interacting males. Pulse rate of calls primarily varied with water temperature. Call rate and call duration showed the most variation of the four call properties, but this variation was unrelated to male morphology or social interactions. Females varied in three aspects of mating behaviour: two measures of pair formation and their preference for dominant frequency of male calls. The body size of paired males varied between females both in pairings initiated by either sex and in pairings initiated only by females. Males chosen by females were usually larger than average, although age and prior breeding experience of females did not affect mate choice. Playback experiments indicated that female preference for calls of low dominant frequency depended on the temporal patterning of alternative calls presented. Each of the four male vocal properties showed significant repeatability, but only one of the three aspects of female mating behaviour was repeatable. We discuss how different degrees of repeatability in sexual traits of males and females may influence the action and detection of sexual selection in this and other species. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Proximate Mechanisms Of Sexual Selection In Wood Frogs

Observations and several types of field experiments on the mating behavior of wood frogs have revealed the proximate mechanisms for a size‐related reproductive advantage in both males and females. For females, larger individuals produce larger clutches; for males, larger individuals can better remain clasped to females when contested by rival males and can better depose males clasped to other females. No results obtained support of the existence of mate choice in either males or females.

THE INFLUENCE OF SEXUAL SELECTION AND INTERSPECIFIC COMPETITION ON MOCKINGBIRD SONG (MIMUS POLYGLOTTOS)

Versatility and individual variation in the song of mockingbirds, Mimus polyglottos, provide an excellent system for studying effects of natural selection. My observations focus on three aspects of mockingbird song: repertoire size, song diversity, and interspecific mimicry. Two hypotheses, not mutually exclusive, are tested: 1) Sexual selection influences repertoire size; males with larger repertoires have an advantage in acquiring territories and attracting mates. Repertoire size is enlarged through interspecific imitations. 2) Interspecific mimicry of territorial songs enables males to exclude close competitors from their territories. By reducing interspecific competition, better mimics have a higher reproductive success. The major selective force influencing bird song is sexual selection. Darwin (1871) postulated two aspects of sexual selection: intrasexual selection for those traits that function in acquisition and maintenance of a breeding territory, and intersexual selection for those traits that affect mate selection. Often intrasexually selected traits also influence mate selection. Primary (or advertizing) song functions both in intraspecific territorial defense and mate attraction. However, intrasexual and intersexual selective pressures are treated separately here to aid in discussion. Intrasexual competition for breeding territories is based on relatively qualitative traits such as aggression as well as quantitative attributes such as plumage characteristics and song. Traits that intimidate rivals will be selected. The

Mate choice and mate competition influence male body size in Japanese medaka

A sexual size dimorphism usually occurs when size-dependent reproductive advantages exist in only one sex. Studies on Japanese medaka, Oryzias latipes, have demonstrated reproductive size advantages in females but not in males, even though males and females are similar in body size. We conducted mate-choice and mate-copying tests in which a female could first associate with, then mate with, either a large (>/=1 sd+X standard length) or a small male (</=1 sd-X standard length). Large males obtained a mating advantage in both tests, and both mate choice and mate competition influenced their mating success. In the majority of trials, females associated with large males. Association preferences of females corresponded to their mating preferences when mate competition between males was weak; however, when mate competition was strong, large males obtained almost every mating regardless of female association preference. Preference for large males may provide females with a reproductive advantage if males mate multiply because small males become sperm-depleted sooner than large males. We found no indication that females copied the mating decisions of other females. Repeatability of female mating preference was low, not because females mated at random with respect to male size, but because most females consistently preferred large males. We also conducted mating tests at four density levels and found that large males maintained their mating advantage relative to small males at all densities. Thus, male and female medaka may be similar in body size because large size provides a fecundity advantage for females, as demonstrated in previous studies, and large size provides a mating advantage for males, as demonstrated in our study. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Assessment of possible ecological risks and hazards of transgenic fish with implications for other sexually reproducing organisms

Transgenic technology is developing rapidly; however, consumers and environmentalists remain wary of its safety for use in agriculture. Research is needed to ensure the safe use of transgenic technology and thus increase consumer confidence. This goal is best accomplished by using a thorough, unbiased examination of risks associated with agricultural biotechnology. In this paper, we review discussion on risk and extend our approach to predict risk. We also distinguish between the risk and hazard of transgenic organisms in natural environments. We define transgene risk as the probability a transgene will spread into natural conspecific populations and define hazard as the probability of species extinction, displacement, or ecosystem disruption given that the transgene has spread. Our methods primarily address risk relative to two types of hazards: extinction which has a high hazard, and invasion which has an unknown level of hazard, similar to that of an introduced exotic species. Our method of risk assessment is unique in that we concentrate on the six major fitness components of an organisms life cycle to determine if transgenic individuals differ in survival or reproductive capacity from wild type. Our approach then combines estimates of the net fitness parameters into a mathematical model to determine the fate of the transgene and the affected wild population. We also review aspects of fish ecology and behavior that contribute to risk and examine combinations of net fitness parameters which can lead to invasion and extinction hazards. We describe three new ways that a transgene could result in an extinction hazard: (1) when the transgene increases male mating success but reduces daily adult viability, (2) when the transgene increases adult viability but reduces male fertility, and (3) when the transgene increases both male mating success and adult viability but reduces male fertility. The last scenario is predicted to cause rapid extinction, thus it poses an extreme risk. Although we limit our discussion to aquacultural applications, our methods can easily be adapted to other sexually reproducing organisms with suitable adjustments of terminology.

Fitness Components and Ecological Risk of Transgenic Release: A Model Using Japanese Medaka (Oryzias latipes)

Any release of transgenic organisms into nature is a concern because ecological relationships between genetically engineered organisms and other organisms (including their wild‐type conspecifics) are unknown. To address this concern, we developed a method to evaluate risk in which we input estimates of fitness parameters from a founder population into a recurrence model to predict changes in transgene frequency after a simulated transgenic release. With this method, we grouped various aspects of an organism’s life cycle into six net fitness components: juvenile viability, adult viability, age at sexual maturity, female fecundity, male fertility, and mating advantage. We estimated these components for wild‐type and transgenic individuals using the fish, Japanese medaka (Oryzias latipes). We generalized our model’s predictions using various combinations of fitness component values in addition to our experimentally derived estimates. Our model predicted that, for a wide range of parameter values, transgenes could spread in populations despite high juvenile viability costs if transgenes also have sufficiently high positive effects on other fitness components. Sensitivity analyses indicated that transgene effects on age at sexual maturity should have the greatest impact on transgene frequency, followed by juvenile viability, mating advantage, female fecundity, and male fertility, with changes in adult viability, resulting in the least impact.

Differential effects of mate competition and mate choice on eastern tiger salamanders

Male tiger salamanders, Ambystoma tigrinum tigrinumare slightly larger in body size and have considerably higher and longer tails than females. To determine how these dimorphic traits affected reproductive performance and success, we conducted breeding trials using 12 males and six females per trial and monitored male-female and male-male interactions. Larger males had an advantage in most aspects of mate competition investigated. Males with higher tails had no advantage in either mate competition or mate choice. Males with longer tails also had no advantage in mate competition but were preferred as mates by females. Larger males interrupted courting males more often than smaller males did. The form of male-male interference was conditional on body size and not on either tail dimension. If the intruder was larger than the courting male, it would shove the female away from the courting male and initiate courtship; if the intruder was smaller, it adopted a female mimicry tactic in which it positioned itself between the courting male and female and performed female behaviours to the courting male while simultaneously courting the female. Our trials indicated that the two components of sexual selection may influence the evolution of different male morphological traits in tiger salamanders. Mate competition may favour increased male body length; mate choice may select for greater male tail length.

ON ALTERNATIVE REPRODUCTIVE TACTICS IN ANURANS: DYNAMIC GAMES WITH DENSITY AND FREQUENCY DEPENDENCE

Intense mating competition and mate choice has favored the evolution of alternative mating tactics in many species. We developed a stochastic dynamic game to evaluate the choice among three alternative mating tactics by male anurans: call, satellite, or leave the chorus to forage or hide. The strategies were assumed to differ in mating success (call > satellite forage = hide), predation risk (call > satellite > forage > hide), and energetic expense (call > satellite = hide > forage). Six major predictions were derived. First, in relatively stable environments, males should enter a chorus in a synchronous pulsed pattern, with first-year males primarily choosing to be satellites on older calling males. The pulsed pattern is caused by the relatively high energetic cost of calling and the dependence of female arrival rates on chorus size (which promotes synchronization). In unstable environments, energetic constraints are reduced because favorable conditions are rare, so most males should call on favorable nights. Under all conditions, energy-constrained males should forage. Second, with an accelerating relationship between female arrival rate (measured per calling male) and chorus size, first-year males in a stable environment should drop out of choruses before the end of the breeding season because the payoff to attendance decreases as the population declines; with a decelerating function, first-year males should remain the entire year. In an unstable environment, males should enter a chorus on favorable days, irrespective of the shape of the female arrival function. Third, variation in the form of predation risk with respect to chorus size has little or no effect on mating patterns; however, high mean predation risk should increase the frequency of satellites in the chorus. Fourth, conflicts can occur between old males that are selected to call and young males that are selected to act as satellites. Under some conditions, the only resolution to this conflict is for all males to abandon the chorus. Fifth, if there are differences among first-year males in when they come into breeding condition, the few males that mature early should be satellites. Later in the season, first-year males should tend to call. This prediction is unaffected by density dependence in predation risk, but it does not hold when female arrival rate is density independent. Finally, first-year males should call more when their mating success decreases with an increase in the number of calling older males in comparison with conditions in which mating success is a fixed fraction of the mating success of older males. We show that the properties of a number of anuran species make them amenable to test these predictions.

SEXUAL SELECTION IN AMERICAN TOADS: A TEST OF A GOOD-GENES HYPOTHESIS

Adaptive mate choice in species lacking male resource control and/or paternal care might be maintained by selection because preferred males sire genetically superior offspring. For such a process to occur, some male phenotypic trait(s) must both reliably indicate male genetic quality and influence the pattern of mate choice by females. In American toads, Bufo americanus, male body length has been documented to influence female mating patterns: females usually mate with males that are larger than average. However, the relationship between male size and male genetic quality is unknown. We conducted a controlled breeding experiment using 48 sires and 19 dams to determine if larger males sire offspring with superior larval performance characteristics (greater survival to metamorphosis, larger mass at metamorphosis, and earlier metamorphosis). We also aged each sire to test the hypothesis that older males are, on average, genetically superior to younger males. We crossed each female with three sires representing three body size categories (mean and 1 SD ± mean snout‐ischium length). Hatchlings (500 from each cross) were reared to metamorphosis in seminatural ponds in the field. Metamorph weight (log transformed) and age at metamorphosis showed significant heritability and were genetically correlated with each other. Hence, sires differed in genetic quality. However, none of the three measures of offspring performance was correlated with sire body size or age. Thus, we obtained no support for the prediction that sire body size or age is related to genetic quality.

Sexual selection on male body size and mating behaviour in American toads, Bufo americanus

Abstract Mating behaviour of B. americanus was observed from 1985 to 1987. The population contained 38–45 males and 11–26 females, depending on the year. The breeding season of this ‘explosive breeder’ usually encompassed less than 48 h. Male reproductive success varied from zero to two matings per season and zero to an estimated 15 126 zygotes per season. All females mated once per season and variation in their zygote production was estimated to be 4017–11 624 zygotes per season. Body length explained 76% of the variation in zygote production of females. However, male body length was only weakly correlated with mating success in two of three seasons, and with zygote production in one of three seasons. Male arm length was predicted to correlate with male mating success because longer arms should facilitate remaining clasped to females when challenged by rival males. However, mating males did not differ from non-mating males in arm length, and the relationship between arm length and body length was the same for the sexes. Various male behaviours were measured using focal-animal sampling but only call rate correlated with male mating success. The pattern of size dimorphism (females larger than males) is consistent with the observed sex-specific relationships between reproductive success and body size.