Association patterns of sailfin mollies (Poecilia latipinna): alternative hypotheses

Individuals may associate with each other due to a variety of selective forces, such as intra- and intersexual selection, and conspecific recognition. Previous studies have concluded that mate choice governs association behavior in polygynous species of fish. I examined whether mate choice underlies the preference for larger individuals by examining preference for association (time spent in proximity to a fish) not only between opposite-sex individuals but also between same-sex individuals of the live-bearing sailfin molly (Poecilia latipinna). Males and females from three size classes were tested with a large and a small object fish of the same and opposite sex. Females preferred to associate with larger over smaller males. Males also preferred to associate with larger over smaller females, as expected. The same female and male test fish also preferred to associate with larger over smaller fish of the same sex. Moreover, females demonstrated no significant difference in their strength of preference (large–small) when offered males or females. The same held true for males. When males and females were subsequently tested with one large male and one large female, females tended to prefer large males while males showed no significant preference for association based on sex. In another experiment, females were tested with a large female and a small male, and significantly preferred the former. These findings suggest that association patterns may have arisen under a variety of conditions, such as predation pressures, shoaling behavior, and associative preference behavior. The assumption that association behavior is a uniformly sufficient predictor of mate choice in fish needs to be re-examined for P. latipinna and other species. Received: 6 November 1998 / Received in revised form: 12 May 1999 / Accepted: 12 May 1999     

Current analyses do not resolve whether extra-pair paternity is male or female driven

Recent studies question the importance of indirect genetic effects in explaining female benefits of extra-pair matings in socially monogamous species. Compiling data on 14 wild bird species, Arnqvist and Kirkpatrick (Am Nat 165:S26–S37, 2005) estimated the average direct cost in terms or reduced parental care to be an order of magnitude larger than the potential effect of genetic benefits. This study has sparked a debate regarding potential confounding factors but no consensus appears to have been reached. Here we focus on the implicit assumption that all individuals face the same selective pressures and argue that this assumption is probably too strong in most cases. Using a theoretical model we show that when the amount of resources that a male provides depends on territory quality, his physical condition or prospects for alternative breeding opportunities, a female may respond to such differences by altering her mating behaviour. Such confounding factors may lead to direct fitness effects that result in negative correlations between paternal care and paternity even if females that produce extra-pair young experience a net benefit. Negative correlations can also result when males forcefully seek copulations and females resist them. We discuss the studies included in the analysis in this light, and conclude that current analyses on the net selective pressures remain uninformative. In addition to considering average effects across individuals and species we suggest giving attention to individual differences and the influence of ecological factors such as territory quality and predation pressures on female mating behaviour.     

The tactics of mutual mate choice and competitive search

A model of mutual mate choice is described, formulated as a dynamic game, which yields predictions about mating behaviour under the influence of time constraints, choice costs and competition for mates. These variables were examined because they may result in a change in the distribution of qualities among unmated individuals of both sexes over the course of the breeding season. The model predicts that mutual choice gives rise to assortative mating, although high costs of choice and/or inaccurate assessment both lead to lower overall correlations between the qualities (or the attractiveness) of mates. When all individuals are present from the start of the breeding season, the correlation between the qualities of individuals pairing at a given time declines throughout the season, so that mates are more closely matched among individuals who pair early than among those who pair late (and extra-pair copulation may thus be more common among the latter). Delayed arrival of lower-quality individuals may, however, lead to an increase in this correlation with time during the early part of the season. The mean quality of unmated males and females declines over time, because more attractive individuals tend to mate sooner. As a result of this decline, and because of time constraints, superior individuals become less choosy as the season progresses. If choice is costly, however, then inferior individuals become more selective with time during the early part of the season, and the level of choosiness peaks later for such individuals. Received: 28 March 1996 / Accepted after revision: 21 September 1996     

Time constraints and multiple choice criteria in the sampling behaviour and mate choice of the fiddler crab,Uca annulipes

Active female sampling occurs in the fiddler crab Uca annulipes. Females sample the burrows of several males before remaining to mate in the burrow of the chosen partner. Females time larval release to coincide with the following nocturnal spring tide and must therefore leave sufficient time for embryonic development after mating. Here we show how this temporal constraint on search time affects female choosiness. We found that, at the start of the sampling period (when time constraints are minimal), females selectively sample the larger males in the population. Towards the end of the sampling period (when the temporal constraints increase the costs of sampling), females are less selective. Furthermore, we suggest that the number of males sampled (and other indices of ‘‘sampling effort’’) may not be reliable indicators of female choosiness and may not reflect the strength of female mating preferences under certain conditions. Burrow quality also emerged as an important criterion in final mate choice. Burrow structure potentially influences reproductive success, and mate acceptance based on burrow structure appears to involve a relatively invariant threshold criterion. Since there is no relationship between male size and burrow quality, females are using at least two independent criteria when choosing potential mates. We envisage mate choice as a two-stage process. First, females select which males to sample based on male size. They then decide whether or not to mate with a male based on burrow features. This sampling process explains how two unrelated variables can both predict male mating success. Received: 23 March 1995/Accepted after revision: 14 January 1996     

Female mate choice in the zebra finch — the effect of male beak colour and male song

Studies of female mate preference in zebra finches (Taenopygia guttata) have shown that male beak colour and song rate are important. However, the two characters are correlated. Here the effect of beak colour and song rate on female choice are examined independently. In mate choice tests involving two males, beak colour was manipulated artificially using nail varnish. The results showed that females showed a significant preference for males with a high song rate, but not with a red beak. Females did not prefer males with a red beak if song rate was low and females preferred males with orange beaks who expressed a high song rate. Female preference for males with red beaks was not found when beak and song characters were no longer correlated.     

Measuring mate choice using correlation: the effect of female sampling behaviour

For intersexual selection to occur, it is necessary that females choose between males. It is now well appreciated that constraints exist, which preclude females sampling all the available males in a population. These constraints are likely to have caused the evolution of sampling rules (such as the “best-of-n” rule) by which females sample males. Here we investigate the impact of female subsampling of the male population, not on the evolution of sampling behaviour, but on the population-level correlation between a male trait and currencies such as reproductive success. This study is important as it illustrates when population-level correlations can be safely used to infer the presence and strength of sexual selection in the field. We find that the correlation between a male trait and a mate choice variable rises steeply as the number of males sampled by each female increases, flattening above seven to ten males sampled. This shape is found to be remarkably robust, and little affected by, for example, the mate choice variable used, by noise in assessment, by sampling behaviour depending on female quality, or by population size. The only variable found to have a large impact is male clumping according to their “quality”. If females are sampling about four males, the maximum correlation that can be found at the population level is in the range 0.4–0.6, perhaps as little as 0.1 if males are strongly clumped. A recent review of the literature suggests that four is the average number of males that females sample. Thus, the absence of a strong correlation cannot by itself be used to infer that sexual selection is weak, as it may be due to females sampling few males. Received: 18 May 1998 / Accepted after revision: 18 July 1998     

Supply and demand: causes and consequences of assortative mating in common terns <Emphasis Type="Italic">Sterna hirundo</Emphasis>

Although assortative mating is widespread among long-lived monogamous birds, the underlying mechanisms still remain unclear in many species. In this study, we analysed assortative mating by age and body mass in the common tern Sterna hirundo with special regard to mate choice decisions of newly paired birds. To assess whether assortative mating by age is an active decision involving either homotypic or directional preferences or rather a passive process due to restricted availability of potential mates, we analysed the influence of age-dependent arrival date and cohort size. Furthermore, we looked for direct benefits in terms of reproductive success. Common terns mated assortatively by age and arrival date but not by body mass at arrival. Assortative mating by age was age dependent and was detected mainly in younger birds, whereas birds older than 8 years rather re-paired with younger ones. The availability of same-aged mates was restricted by cohort size and arrival date. We found no general evidence for better reproductive performance of same-aged pairs. Instead, relative reproductive success was related to own age and the relative age of mate: It should be advantageous for any bird to acquire an old mate; hence, assortative mating by age seems to be beneficial only for old terns but not for young ones. Age-assortative mating in common terns occurred by both passive and active processes, which are not mutually exclusive. Our results do not indicate a homotypic but a directional preference and support the theory that high-quality (older/experienced) individuals tend to mate assortatively if same-aged mates are available, which leaves low-quality (young/inexperienced) individuals to mate among themselves.     

Sexual conflict over fertilizations: female bluethroats escape male paternity guards

Extra-pair copulations create a potential for sexual conflict in pair-bonding birds. Here we report an experimental study of the bluethroat, Luscinia s. svecica, in which the throat ornament of males was blackened with Nyanzol D in order to reduce their sexual attractiveness and thus increase the sexual conflict over fertilizations. In an earlier study, we showed that males blackened before pairing had a lower success in attracting social mates than controls, whereas males blackened after pairing guarded their mates more intensely and sang less than controls. Here we add behavioural data from one more year on males blackened after pairing and corroborate our previous finding that the manipulation caused males to guard their mates more intensely and advertise less for additional mates. Blackened males did not suffer more intrusions from neighbouring males than did controls. Paternity analyses of the combined data set, using multilocus DNA fingerprinting and microsatellite typing, revealed that blackened males lost significantly more paternity than controls. There was also a tendency for blackened males to show a lower success in achieving extra-pair fertilizations. These results indicate that females have the upper hand in the sexual conflict over fertilizations, as females paired with unattractive males can achieve more extra-pair paternity despite the greater constraint posed by the intensified mate guarding. Still, within the blackened group, there were some indications that males guarding more intensely and singing less had higher paternity than males guarding less and singing more, suggesting a marginal positive effect of guarding for unattractive males. Male mate guarding must nevertheless be considered a best-of-a-bad-job strategy in this species. Received: 4 December 1997 / Accepted after revision: 14 June 1998     

Male-male association patterns and female proximity in the guppy,Poecilia reticulata

If males differ in their ability to attract potential mates, and are able to perceive such differences, theory predicts they should distribute themselves in a manner that increases their probability of obtaining potential matings. The relationship between male-male association patterns and the proximity of females in social groups, however, remains virtually unexplored. Experimental analysis of this relationship in the guppy, Poecilia reticulata, demonstrates that in preference tests males showed a strong tendency to associate with other males that were further away from potential mates than they were themselves. Male guppies pursue a behavioral strategy that involves categorizing other males based on their proximity (and possibly relative attractiveness) to females, remembering the identity of such individuals, and using this information when choosing between other males as associates. Such a strategy may increase a male's chances of being the individual chosen by a female assessing nearby males.     

Sperm allocation in an uncertain world

Models of optimal sperm allocation are constructed using stochastic dynamic programming techniques, yielding predictions about sperm allocation when males mate sequentially, breeding time is finite, sperm stores are limited and females vary in quality. The models suggest that uncertainty of future reproductive opportunities should favour withholding of sperm, so that males tend to allocate less (for a given level of stored sperm) earlier in the breeding season. This effect is more pronounced the greater the variance in female quality. We also show that while allocation will be influenced by mate value, it is not necessarily optimal to allocate preferentially to high-quality females, since the benefits of a higher-quality mate may be offset by increased risk of rejection of sperm or higher sperm competition. The relationship between mate quality and level of allocation will depend strongly on the amount of remaining stored sperm, with males whose supplies are depleted being more likely to favour lower-quality partners. Received: 12 September 1997 / Accepted after revision: 28 June 1998