Trond Amundsen

Why are female birds ornamented

Sexual selection is now widely accepted as the main evolutionary explanation of extravagant male ornaments. By contrast, ornaments occurring in females have received little attention and often have been considered as nonadaptive, correlated effects of selection on males. However, recent comparative evidence suggests that female ornaments have evolved quite independently of male showiness. Also, new theoretical models predict that both male mate choice and female contest competition will occur under certain circumstances. This is supported by recent experimental studies. Thus, selection acting on females might be a widespread cause of female ornaments.

Male mate choice selects for female coloration in a fish

Although sexual selection theory has proved successful in explaining a wide array of male ornaments, the function of ornaments occurring in females is largely unknown. Traditionally, female ornaments have been considered nonfunctional, being merely a genetically correlated response to selection for male ornamentation. However, this hypothesis is only relevant to species in which the ornament is basically the same in the two sexes. Alternatively, female ornaments may be influenced by selection acting directly on the females, either through female–female competition or male choice. We tested the latter hypothesis in mate-choice experiments with two-spotted gobies (Gobiusculus flavescens). In this small marine fish, females have bright yellow-orange bellies during the breeding season, a conspicuous trait that is not present in males. We conducted two aquarium experiments to test whether males preferred to mate with more colorful females. In the first experiment, males had a choice between two females that varied in natural coloration (and belly roundness). In the second experiment, we manipulated belly coloration and kept roundness constant. Males spent more time with colorful than with drab females in both experiments and also performed far more courtship displays toward colorful females. Our study provides experimental evidence that males prefer ornamented females in a fish that is not sex-role reversed, supporting the hypothesis that female ornamentation is sexually selected.

Ultraviolet colour vision and ornamentation in bluethroats

Many birds see in the ultraviolet (300–400 nm), but there is limited evidence for colour communication (signalling by spectral shape independently of brightness) in this ‘hidden’ waveband. Such data are critical for the understanding of extravagant plumage colours, some of which show considerable UV reflectance. We investigated UV colour vision in female social responses to the male UV/violet ornament in bluethroats, Luscinia s. svecica. In an outdoor aviary at the breeding grounds, 16 females were each presented with a unique pair of males of equal age. In UVR (UV reduction) males, sunblock chemicals reduced only the UV reflectance and thereby the spectral shape (colour) of the throat ornament. In NR (neutral reduction) males, an achromatic pigment in the same base solvent (preen gland fat) was used for a corresponding but uniform brightness reduction. Both colour and brightness effects were invisible to human eyes, and were monitored by spectrometry. In 13 of the 16 trials, the female associated most with the NR male, a preference that implies that UV colour vision is used in mate choice by female bluethroats. Reflectance differences between one–year–old and older males were significant only in UV, suggestive of a UV colour cue in age–related mate preferences.

Unusually dynamic sex roles in a fish

Sex roles are typically thought of as being fixed for a given species. In most animals males compete for females, whereas the females are more reluctant to mate. Therefore sexual selection usually acts most strongly on males. This is explained by males having a higher potential reproductive rate than females, leading to more males being sexually active (a male-biased operational sex ratio). However, what determines sex roles and the strength of sexual selection is a controversial and much debated question. In this large-scale field study, we show a striking temporal plasticity in the mating competition of a fish (two-spotted goby, Gobiusculus flavescens). Over the short breeding season fierce male–male competition and intensive courtship behaviour in males were replaced by female–female competition and actively courting females. Hence, sex role reversal occurred rapidly. This is the first time that a shift in sex roles has been shown in a vertebrate. The shift might be explained by a large decline in male abundance, strongly skewing the sex ratio towards females. Notably, the sex role reversal did not occur at an equal operational sex ratio, contrary to established sex role theory.

On the function of female ornaments: male bluethroats prefer colourful females

Female ornaments in animals with conventional sex roles have traditionally been considered non–functional, being merely a genetically correlated response to selection for male ornamentation. Alternatively, female ornaments may be influenced by selection acting directly on the females, either through female–female competition or male choice. We tested the latter hypothesis in mate choice experiments with bluethroats (Luscinia s. svecica), a passerine bird in which females vary considerably in coloration of an ornamental throat patch. In outdoor aviaries placed in prime breeding habitat, males were allowed to choose between a colourful and a drab female. We found that males associated more with, and performed more sexual behaviours towards, colourful females. Female coloration was not age–related, but correlated significantly with body mass and tarsus length. Thus, we have demonstrated both a male preference for female ornamentation, and a relationship between ornament expression and female body size, which may be indicative of quality. Our results refute the correlated response hypothesis and support the hypothesis that female ornamentation is sexually selected.

Costs and benefits of hatching asynchrony in blue tits Parus caeruleus

We studied the significance of hatching asynchrony in blue tits Parus caeruleus during a 4-year period at Oslo, Norway, by comparing breeding success of broods manipulated to hatch over a shorter or longer period than average. Nestling mortality was high and mainly caused by starvation. Mean body mass of fledglings on day 14 was significantly higher for asynchronous than for synchronous broods. No significant difference was found in the mean number of young fledged between the treatment groups, nor in the number of offspring recruited into the local breeding population, even in a year with heavy nestling mortality. However, the number of recruits was few. The results are consistent with the view that asynchronous hatching ensures high quality of some offspring (the offspring quality assurance hypothesis). This benefit is achieved even when no brood reduction occurs. We suggest a potential cost to hatching asynchrony (the diminishing return hypothesis): in asynchronous broods, early hatched offspring may require so much food that their own subsequent survival is reduced or at least no further improved. Resources may therefore be wasted on first hatched offspring that might have improved the growth of younger siblings. Female parents had lower post-breeding survival with asynchronous than with synchronous hatching; the opposite result was found for male parents. We propose the exploitation of mate hypothesis to explain this result. The hypothesis presumes there is sexual conflict over the amount of parental investment in current versus future reproduction, and over the investment in particular offspring within the brood. We suggest that with asynchronous hatching, females have to invest more to keep less competitive, late-hatching nestlings alive.

Effects of egg size and parental quality on early nestling growth: An experiment with the Antarctic petrel

A large number of studies have reported a positive relationship between the egg size of birds and the subsequent growth and/or survival of nestlings, but such effects may partly be due to confounding variables, e.g. parental quality. In order to evaluate the potential effects of egg size, and of parental quality, on early nestling growth in the Antarctic petrel, we performed an experiment in which eggs of different size were swapped between nests. From a sample of 300 nests with eggs of known size, we selected eggs belonging to the lower quartile (small eggs), and those belonging to the upper quartile (large eggs), with respect to volume. Half of the small eggs were exchanged with small eggs from other nests, and the other half with large eggs. A similar procedure was used for large eggs. Growth and survival of the nestlings were recorded until 12 days old. Hatching success was positively related to egg size. Egg size influenced nestling body mass until the age of 3 days, and tarsus length was affected until 12 days old. However, these effects were not due to an effect of egg size on growth rates, but reflected instead the influence of egg size on hatchling size. In contrast to most previous studies, we found no effect of parental quality (as reflected in the size of own eggs) on foster nestling size or growth until 12 days old. This could be because egg size does not reliably reflect parental quality in the species, or because parental effects become evident only at later nestling stages. We discuss why egg size variation is maintained in this and other species where egg size influences parental fitness through the survival of eggs or nestlings.

Seasonal change in female choice for male size in the two-spotted goby

Mate choice may vary according to various factors, such as mate availability and variation in mate quality. We tested seasonal changes in female choice regarding male size in a marine fish, the two-spotted goby, Gobiusculus flavescens. In this species, males occupy nests in which females lay their eggs and males thereafter provide exclusive parental care of the eggs until hatching. Females are predicted to prefer large males if, for example, they are better at competing for and defending nests. In this species, the opportunity for females to be selective should decline over the breeding season as the availability of males decreases. We therefore predicted a reduction in female choosiness with season. We conducted mate choice tests, during the early (May) and later (June) part of the breeding season, in which females were allowed to choose between males differing in body length. Females chose large males early in the season, but became unselective with respect to male size later on. This change in mate choice may be caused by a reduction in overall choosiness as a result of the decline in male availability with season. Alternatively, the change could be caused by a switch in choice cues used by females if factors other than male size become more important for female reproductive success later in the breeding season. Further studies are needed to test between these alternative explanations.

Does female plumage coloration signal parental quality? A male removal experiment with the bluethroat (Luscinia s. svecica)

Abstract Females in several sexually dimorphic species with conventional sex roles possess ornamental traits that resemble those found in males. The evolution of such traits, however, is still poorly understood. Bluethroats (Luscinia s. svecica) are socially monogamous, sexually dichromatic passerine birds, in which female throat patch coloration varies from near absence to near full expression of male-like coloration. A recent study, demonstrating that male bluethroats prefer colourful females, suggests that female coloration is subject to sexual selection through male choice. However, the benefits males may gain from mating with colourful females have not yet been identified. In this study we tested the hypothesis that female coloration signals parental quality (the good-parent hypothesis). During the course of the same day, we recorded female care both in the presence and the absence of the male mate. The latter was done to eliminate the confounding effect of variable male care by removing the male temporarily. Female coloration did not correlate with female feeding rates either in the presence or in the absence of the male. Female feeding rates in the absence and the presence of the male were positively, although weakly, correlated. Female coloration did not correlate with female ability to compensate for the loss of male care, or with the change in brood mass during male removal. Therefore, there is no evidence for the good-parent hypothesis to explain female plumage coloration in bluethroats.

Male preference for colourful females affected by male size in a marine fish

It is becoming increasingly clear that mate preferences are not static, but can vary as a function of ecological conditions and the state of the choosing individual. This applies not only to females, the sex that has usually been the subject in research on mate preferences, but also to males. Under certain conditions, males should be selective in their choice of breeding partner. In the two-spotted goby, Gobiusculus flavescens, a small marine fish, breeding females develop conspicuous yellow-orange bellies, which they actively display to males during courtship. We have recently shown that males prefer more colourful females as mates. In this study, we test if the size of a male affects his preference for colourful females. Using three-compartment mate-choice aquaria, we recorded the interest shown by a male in two females differing in coloration but similar in size. Large and small males were equally eager to court females, but only large males showed a greater interest in the more colourful females. We suggest that small males are unselective because they usually obtain few mating opportunities, as a result of being unsuccessful in mate attraction or male contest competition. This study provides the first demonstration that the size of a male affects his preference for female colour.