Mate choice by male sticklebacks, Gasterosteus aculeatus

Territorial male threespine sticklebacks, Gasterosteus aculeatus, were presented simultaneously with two gravid female stickleback dummies that differed only in degree of abdominal distention. Males directed most (approximately 60%) of their courtship behaviour (zigzags and leads) to the more distended dummy. However, when presented with the same two dummies consecutively, males directed similar amounts of courtship to each. Males probably prefer the more distended females because distention indicates greater fecundity or spawning readiness.     

Mate choice by hen Red Grouse Lagopus lagopus with an excess of cocks—role of territory size and food quality

In a breeding population of Red Grouse the ratio of cocks to hens was about 2:1. Cocks with hens had bigger territories and richer feeding sites than unmated cocks. Despite this, the size of a cock's territory was inversely related to the level of nitrogen in his food. The idea that this was because cocks defend a territory just big enough to meet the pair's food requirements seemed inapplicable, since territories were much bigger than this. In addition, it was inferred that hens took longer to choose a cock on a part of the area that had relatively poor feeding sites. Whether hens chose the cock or his territory was uncertain, but both probably played a part.     

Female marsh wrens do not provide evidence of anatomical specializations of song nuclei for perception of male song

In songbirds the forebrain nuclei HVC (high vocal center) and RA (robust nucleus of the archistriatum) are larger in individuals or species that produce larger song repertoires, but the extent to which the size of these nuclei reflects a need for either producing or perceiving large repertoires is unknown. We, therefore, tested the hypothesis that species differences in the size of song nuclei reflect a commitment of “brain space” to the perceptual processing of conspecific song. The two species of marsh wren (Cistothorus palustris western and eastern) provide a good test case. Western males produce larger song repertoires, and have larger HVC and RA than do eastern males. Female marsh wrens do not sing, and if they use their song nuclei to assess conspecific male song repertoires, then we predicted that measurable cellular and nuclear parameters of HVC and RA would be greater in western than eastern female wrens. For males we confirmed that the volumes of HVC and RA, and cellular parameters of HVC, are greater in western than in eastern birds. These nuclei were also considerably larger in males than in conspecific females. Western and eastern female wrens, however, did not differ in any measured parameters of HVC or RA. Females of these wren species thus do not provide any direct evidence of anatomical specializations of song nuclei for the perceptual processing of conspecific male song. 1994 John Wiley & Sons, Inc.     

Mate choice for genetic quality when environments vary: suggestions for empirical progress

Mate choice for good-genes remains one of the most controversial evolutionary processes ever proposed. This is partly because strong directional choice should theoretically deplete the genetic variation that explains the evolution of this type of female mating preference (the so-called lek paradox). Moreover, good-genes benefits are generally assumed to be too small to outweigh opposing direct selection on females. Here, we review recent progress in the study of mate choice for genetic quality, focussing particularly on the potential for genotype by environment interactions (GEIs) to rescue additive genetic variation for quality, and thereby resolve the lek paradox. We raise five questions that we think will stimulate empirical progress in this field, and suggest directions for research in each area: (1) How is condition-dependence affected by environmental variation? (2) How important are GEIs for maintaining additive genetic variance in condition? (3) How much do GEIs reduce the signalling value of male condition? (4) How does GEI affect the multivariate version of the lek paradox? (5) Have mating biases for high-condition males evolved because of indirect benefits?     

Stylish lengths: Mate choice in flowers

The styles of flowers may represent an arena for pollen competition in the race to fertilize ovules. Accordingly, selection should favour a longer ‘race’ to better discriminate among variable pollen by increasing style length. Sampling across a taxonomically diverse range of wild and outcrossed species, we found that the distribution of style lengths within plants were skewed towards longer styles, as predicted. In self-pollinated domesticated species, where discrimination among pollen is less important, we found no such pattern. We conclude that style length is under directional selection towards longer styles as a mechanism for mate choice among pollen of variable quality.     

Red coloration of male northern cardinals correlates with mate quality and territory quality

I investigated how mate quality and territory quality influencean extravagant ornament in a socially monogamous species thatdefends multipurpose territories. Northern cardinals (Cardinaliscardinalis) are a highly dichromatic, socially monogamous species,and males are a brilliant red. I conducted a 3-year field studyof northern cardinals and found that redder males produced moreoffspring in a breeding season. Two selective factors mediatedthis fitness gain. Redder males were paired with earlier breedingfemales, an established measure of mate quality in birds. Second,redder males obtained territories of higher quality, as measuredby vegetation density. Interactions among these factors werealso important in explaining variance in male reproductive success.Multivariate analysis indicated that earlier breeding increasedreproductive success independent of territory quality. In turn,territory quality contributed to male reproductive success throughits effect on nest survival and possibly through its role in attractingan earlier breeding female.     

Female mate choice based on territory quality in barn swallows

Female mate choice based on territory quality is difficult to study because territories often contain many resources, which are difficult to quantify. Here, using the barn swallow (Hirundo rustica gutturalis) breeding at an outdoor breeding site in Japan, where each male defends only a small territory containing old nests, we studied whether females choose social mates based on territory quality. Since the territories of this species contain few other resources, territory quality can easily be assessed by quantifying old nests in the territory. We made the following four observations: (1) male swallows displayed old nests in their territories to females; (2) the old nests used for the first clutch were less broken than the other old nests within the same territory; (3) territory quality, defined by the number of old nests weighted by the intactness of each old nest, predicted the productivity of the territory; and (4) males with better territories paired with females earlier, and hence bred earlier, than those with inferior territories. The relationships remained significant even after controlling for male morphological traits. Based on these results, we can infer that female swallows choose their mates based, in part, on territory quality.     

Effects of location in territory and distance from neighbours on the use of song repertoires by Carolina wrens

By capturing territorial Carolina wrens (Thryothorus ludovicianus) and housing them in portable cages on their territories, I could control each one's location in its territory and distance from neighbours in experiments on the use of song repertoires. Experiment 1 demonstrated that these wrens sang more songs in the centre of their territories than at the edge, but that they did not use more song types or different song types at the centre than at the edge. In experiment 2, in which I played tape-recorded songs at two distances from wrens caged in the centres of their territories, birds responded more strongly to songs at 25m, simulating an intruder, than to songs at 165 m, simulating a territorial neighbour. Birds also switched more frequently between song types and sang more song types per 100 songs in response to the nearer playback. Experiment 3 compared captive wrens 140 m, 80 m, 20 m, or 0 m apart on adjacent territories. As the distance between neighbours decreased, birds sang less, but also switched more frequently between song types, used more song types per 100 songs, and matched songs with neighbours more frequently. There were no differences in the kinds of song types sung at different distances from neighbours. A comparison of the results from experiments 1 and 3 confirms that the use of song repertoires is influenced by distance from conspecifics and not by location in the territory.     

Mate discrimination by females in the burying beetle Nicrophorus orbicollis: the influence of male size on attractiveness to females

1. Female burying beetles behave differently towards males of different sizes, avoiding mating with large males that are not defending resources but mating with small males regardless of the presence of resources. Females of the burying beetle Nicrophorus orbicollis were therefore examined to determine whether they discriminate among males using only pheromonal signals. The influence of female size on its own mate choice was also examined. 2. Females do use male pheromonal signals to discriminate among males and these signals do appear to convey information about male body size to females. Overall, females were more likely to be attracted to larger males than to smaller males. 3. Female choice of a male was influenced by both the female's own body size and the size of the female relative to the size of the two males available to it. 4. While there is an overall mating advantage for larger males, resulting from female preferences based on odour cues, smaller males are also attractive to some females under some circumstances. 5. It is argued that there are different costs and benefits of mating with different sized males, leading to the evolution of context‐dependent mate choice for females and the need to be able to discriminate males of different sizes from a distance.     

Female choice in the sedge warbler Acrocephalus schoenobaenus: multiple cues from song and territory quality

Recent models of animal signalling emphasize the evolution of complex displays containing ''multiple messages''. A variety of potential cues used in female choice were investigated during a three-year field study of the sedge warbler, Acrocephalus schoenobaenus. Twelve possible cues were investigated, and three were found to have a significant influence upon pairing date. Two were different measures of song (repertoire size and song flighting) and one a measure of territory (territory size). Repertoire and territory size had a significant influence on pairing date in all three years, and song-flighting in two. The three cues were not intercorrelated and so had independent effects upon pairing date. We suggest that females select males upon multiple cues as these reflect different aspects of male and territory quality.     

Intraspecific egg destruction in marsh wrens: a study of mechanisms preventing filial ovicide

Marsh wrens, Cistothorus palustris, attack the nests of other birds, including conspecifics. The tendency of wrens to attack conspecific nests raises the question: what prevents breeding individuals from destroying their own offspring? In this study the mechanisms that prevent filial ovicide (i.e. the destruction of eggs by parents) are examined. The results suggest that male filial ovicide is prevented by the presence of a female in the nest area, which presumably inhibits the male's ovicidal tendencies. In addition, incubating females aggressively exclude males from their breeding area, thereby further reducing the threat to their nest. Female filial ovicide is prevented through the inhibition of intraspecific nest-destroying behaviour in breeding females and through the ability of individual females to recognize their own nest site and nest. Differences between the intraspecific nest-destroying behaviour of male and female marsh wrens may reflect different reproductive strategies of the two sexes.     

Mate choice and uncertainty in the decision process

The behavior of females in search of a mate determines the likelihood that a high quality male is encountered in the search process and alternative search strategies provide different fitness returns to searchers. Models of search behavior are typically formulated on an assumption that the quality of prospective mates is revealed to searchers without error, either directly or by inspection of a perfectly informative phenotypic character. But recent theoretical developments suggest that the relative performance of a search strategy may be sensitive to any uncertainty associated with the to-be-realized fitness benefit of mate choice decisions. Indeed, uncertainty in the decision process is inevitable whenever unobserved male attributes influence the fitness of searchers. In this paper, we derive solutions to the sequential search strategy and the fixed sample search strategy for the general situation in which observed and unobserved male attributes affect the fitness consequences of female mate choice decisions and we determine how the magnitude of various parameters that are influential in the standard models alter these more general solutions. The distribution of unobserved attributes amongst prospective mates determines the uncertainty of mate choice decisions-the reliability of an observed male character as a predictor of male quality-and the realized functional relationship between an observed male character and the fitness return to searchers. The uncertainty of mate choice decisions induced by unobserved male attributes has no influence on the generalized model solutions. Thus, the results of earlier studies of these search models that rely on the use of a perfectly informative male character apply even if an observed male trait does not reveal the quality of prospective mates with certainty. But the solutions are sensitive to any changes of the distribution of unobserved male attributes that alter the realized functional relationship between an observed character and the fitness return to searchers. For example, the standard sequential search model exhibits a reservation property--the acceptability of prospective mates is delimited by a unique threshold criterion--and the existence of this model property under generalized conditions depends critically on the association between the observed and unobserved male characters. In our formulations of the models we assumed that females use a single male character to evaluate the quality of prospective mates, but the model properties generalize to situations in which male quality is evaluated by a direct inspection of multiple male characters.     

Mate choice: been there, done that

Females may benefit from mating with several males, but how can they avoid mating with the same male repeatedly? A new study of crickets has found that females mark their mates to avoid repeat copulations.     

Density-dependence by habitat heterogeneity: individual quality versus territory quality

The aim of this comment is to review the ecological issues concerning the role of individual and habitat heterogeneity as possible mechanisms explaining density-dependent fecundity in animal populations. Our intention is to discuss different approaches to determine whether or not studied populations are subjected to density-dependent processes and which mechanisms are involved. We show that because individual quality (e.g. measured in terms of age of breeding individuals) and territory quality can be correlated, untangling both effects on fecundity is frequently a difficult task. We discuss the misuse of statistical methods and other problems related to the specific characteristics of the studied populations that can have a strong influence on the conclusions reached by researchers working in this field.     

Mate choice and imprinting in birds studied by cross-fostering in the wild

Sexual-selection theories generally assume that mating preferences are heritable traits. However, there is substantial evidence that the rearing environment may be important for the development of mating preferences, indicating that they may be learnt, or modified by experience. The relative importance of such sexual imprinting across species remains largely unexplored. Here, we report results of a large-scale cross-fostering experiment in the wild in which nestling birds were raised by parents of a different species. We show that resulting sexual imprinting may have a negative effect on pairing success in one species (the great tit, Parus major), but not in two other species (the blue tit, P. caeruleus and the pied flycatcher, Ficedula hypoleuca). A remarkable variation thus seems to exist, even between species that are congeneric and have similar breeding ecologies. The cross-fostering resulted in heterospecific pairings between the two tit species (female blue tit breeding with male great tit), which has never, to our knowledge, been previously documented. However, the chicks fledging from these nests were all blue tit.     

Mate choice, monogamy and mutational load

The classical Haldane mutational load for dominant mutations is 1 - e-alpha (where alpha = the mutation rate of the genome). This is independent of the fitness of the mutant heterozygotes. As alpha may be between 0.9 and 1.8 this may be too high for low fecundity species to bear. The load may be greatly reduced for mildly deleterious mutations if female choice (for males with few or no mutations) is present in a polygamous species. It is shown here that if females and males prefer partners with few or no mutations then the load may also be reduced in monogamous species. As in the polygamous model the effect is greatest if mutations are mildly deleterious.     

Mate choice in the face of costly competition

Studies of mate choice commonly ignore variation in preferencesand assume that all individuals should favor the highest-qualitymate available. However, individuals may differ in their matepreferences according to their own age, experience, size, orgenotype. In the present study, we highlight another simplereason why preferences may differ: if there is costly competitionfor mates, the poorest competitors might be better off avoidingthe highest-quality partners and instead targeting low-qualitypartners, so that they minimize the costs they incur. We presenta game-theoretical model of mate choice in which males of differingquality compete for access to females and try to retain themtill the time of mating. Our model predicts that high-qualitymales, who are better competitors, have a preference for thebest females that is typically several times stronger than thatof low-quality males. Early in the competitive period, the lattermay even prefer low-quality females over high-quality females.Thus, variation in competitive ability generates variation inboth the strength and direction of preferences. Differencesin competitive ability result in assortative mating with respectto quality, which is reinforced by variation in preferences.As the time of mating draws near and there is an increased riskof ending up unpaired, all males become indifferent to the qualityof potential mates. Our findings are equally applicable to femalechoice for males, and offer a new explanation for adaptive variationin mating preferences based on differing abilities to cope withthe costs of mate choice.