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Female mate choice in a subsocial beetle: male phenotype correlates with helping potential and offspring survival.

In Parastizopus armaticeps (Tenebrionidae), a nocturnal desert beetle, the males excavate breeding burrows and maintain their moisture level while the females provision the larvae with detritus collected on the surface. The beetles court in small groups on the surface at night after rain. Male size distribution in these groups corresponded to that in the population but more large and fewer small females were present than expected and more large beetles of both sexes bred. Offspring number correlated positively with burrow depth and body length for males but not for females. Since large males dig deeper burrows, which results in higher larval survival rate, females should prefer them. In choice experiments, females selected larger males. Behavioural analyses showed that choice was not dependent on differences in male courtship activity or intermale dominance. When the mass of the smaller male was increased experimentally by a dorsally attached weight, the smaller male was preferred, females estimating male size difference by mass. Partner choice is therefore epigamic for a male phenotypic character which correlates with both parenting ability and greater reproductive success for females. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Female swordtail fish prefer symmetrical sexual signal

It has been proposed that females use the symmetry of secondary sexual traits to differentiate between potential mates. The vertical bars on male swordtail fish function as a signal that attracts females and deters rival males in one swordtail species. In addition, male courtship behaviour of most Xiphophorus species incorporates serial lateral presentations, which provide females with a clear opportunity to assess males for bilateral symmetry. We tested the hypothesis that X. cortezi females prefer males with a symmetrical bar number by determining whether females switched their preference between two males when we switched which male had a symmetrical number of bars. The ability to manipulate the bars without influencing other male traits allowed us to control for male characters correlated with bar symmetry that females might prefer. The degree of asymmetry in bar number we used was within the degree of asymmetry found for this trait in nature. Females switched their preference between a pair of males when we switched which male was symmetrical for bar number. We discuss the possibility that females prefer a symmetrical bar number as well as an alternative hypothesis. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Paternal inheritance of a female moth's mating preference.

Females of the arctiid moth Utetheisa ornatrix mate preferentially with larger males, receiving both direct phenotypic and indirect genetic benefits. Here we demonstrate that the female's mating preference is inherited through the father rather than the mother, indicating that the preference gene or genes lie mostly or exclusively on the Z sex chromosome, which is strictly paternally inherited by daughters. Furthermore, we show that the preferred male trait and the female preference for that trait are correlated, as females with larger fathers have a stronger preference for larger males. These findings are predicted by the protected invasion theory, which asserts that male homogametic sex chromosome systems (ZZ/ZW) found in lepidopterans and birds promote the evolution of exaggerated male traits through sexual selection. Specifically, the theory predicts that, because female preference alleles arising on the Z chromosome are transmitted to all sons that have the father's attractive trait rather than to only a fraction of the sons, such alleles will experience stronger positive selection and be less vulnerable to chance loss than would autosomal alleles.

Alleles↗

Variation in mate choice and mating preferences: a review of causes and consequences.

The aim of this review is to consider variation in mating preferences among females. We define mating preferences as the sensory and behavioural properties that influence the propensity of individuals to mate with certain phenotypes. Two properties of mating preferences can be distinguished: (1) "preference functions'-the order with which an individual ranks prospective mates and (2) "choosiness'-the effort an individual is prepared to invest in mate assessment. Patterns of mate choices can be altered by changing the costs of choosiness without altering the preference function. We discuss why it is important to study variation in female mating behaviour and identify five main areas of interest: Variation in mating preferences and costs of choosiness could (1) influence the rate and direction of evolution by sexual selection, (2) provide information about the evolutionary history of female preferences, (3) help explain inter-specific differences in the evolution of secondary sexual characteristics, (4) provide information about the level of benefits gained from mate choice, (5) provide information about the underlying mechanisms of mate choice. Variation in mate choice could be due to variability in preference functions, degree of choosiness, or both, and may arise due to genetic differences, developmental trajectories or proximate environmental factors. We review the evidence for genetic variation from genetic studies of heritability and also from data on the repeatability of mate-choice decisions (which can provide information about the upper limits to heritability). There can be problems in interpreting patterns of mate choice in terms of variation in mating preferences and we illustrate two main points. First, some factors can lead to mate choice patterns that mimic heritable variation in preferences and secondly other factors may obscure heritable preferences. These factors are divided into three overlapping classes, environmental, social and the effect of the female phenotype. The environmental factors discussed include predation risk and the costs of sampling; the social factors discussed include the effect of male-male interactions as well as female competition. We review the literature which presents data on how females sample males and discuss the number of cues females use. We conclude that sexual-selection studies have paid far less attention to variation among females than to variation among males, and that there is still much to learn about how females choose males and why different females make different choices. We suggest a number of possible lines for future research.

Animal Communication↗

Complex Mhc-based mate choice in a wild passerine.

The extreme polymorphism of the vertebrate major histocompatibility complex (Mhc) is famous for protecting hosts against constantly evolving pathogens. Mate choice is often evoked as a means of maintaining Mhc variability through avoidance of partners with similar Mhc alleles or preference for heterozygotes. Evidence for these two hypotheses mostly comes from studies on humans and laboratory mice. Here, we tested these hypotheses in a wild outbred population of house sparrows (Passer domesticus). Females were not more or less closely related to the males they paired with when considering neutral genetic variation. However, males failed to form breeding pairs when they had too few Mhc alleles and when they were too dissimilar from females at Mhc loci (i.e. had no common alleles). Furthermore, pairs did not form at random as Mhc diversity positively correlated in mating pairs. These results suggest that mate choice evolves in response to (i) benefits in terms of parasite resistance acquired from allelic diversity, and (ii) costs associated with the disruption of co-adapted genes.

Animals↗

Sexual selection promotes hybridization between Pecos pupfish, Cyprinodon pecosensis and sheepshead minnow, C. variegatus.

Rapid and extensive genetic introgression has occurred between Pecos pupfish (Cyprinodon pecosensis) and sheepshead minnow (Cyprinodon variegatus) in the wild. We studied both female mate choice and male-male competition for mates among C. pecosensis, C. variegatus, and their F1 hybrids to determine what role these behaviours played in the formation of the hybrid swarm. Female C. pecosensis preferred male C. variegatus to conspecific males, C. variegatus females displayed no significant preference when given a choice between purebred males, and neither C. pecosensis nor C. variegatus females discriminated against F1 hybrid males. We found no evidence for female olfactory recognition of mates. Male F1 hybrids and C. variegatus were more aggressive than C. pecosensis males, achieving greater reproductive success under two different experimentally-induced mating systems. Hybrids were superior to C. variegatus when only two males competed (dominance interactions), but the two types were competitively equivalent in a territorial mating system. Our results indicate that active inter- and intra-sexual selection contributed to the accelerated hybridization between these two species. By including the possibility that some aspects of a hybridization and introgression event may be under positive selection, researchers may better understand the dynamics that lead to hybrid zone stability or the spread of introgressed genetic material.

Aggression↗

Modes and origins of mechanical and ethological isolation in angiosperms.

Mechanical and ethological isolation between species is widespread in angiosperms with specialized animal-pollinated flowers, being recorded in 29 species groups belonging to 27 genera and 16 families. Mechanical isolation occurs in two forms. (i) The common type, designated the Salvia type, operates when two or more species of flowers are adapted for different groups of pollinators with different body sizes and shapes. (ii) In the Pedicularis type two flower species have the same species of pollinator but pick up pollen from different parts of the pollinator's body. Four forms of ethological isolation are recognized. (i) In the Aquilegia type, which is widespread, ethological isolation is a side effect of mechanical isolation. (ii) The flower-constancy type, as the name suggests, is based on flower-constant foraging behavior. (iii) In the Ophrys type, floral scents attract male bees or wasps and play a role in their mating behavior; different species of flowers, often orchids, have different scents and attract different sets of hymenopteran species. (iv) The monotropy type occurs in plants pollinated by hymenopterans with species-specific or group-specific flower preferences for nutritive purposes (monotropic and oligotropic bees and fig wasps). Three modes of origin of floral isolation are confirmed by evidence: (i) mechanical and ethological isolation arising as a by-product of allopatric speciation, (ii) ethological isolation developing by selection for reproductive isolation per se, and (iii) mechanical isolation arising as a by-product of character displacement. Mode of origin i accounts for the Salvia and Aquilegia types of isolation in nine known species groups and for the Ophrys type in one group. Mode of origin ii accounts for the flower-constancy type of ethological isolation in two species groups. Mode of origin iii explains mechanical isolation in two groups. Sympatric origin of floral isolation by hybrid speciation and by flower constancy has been proposed, but these modes are undocumented and improbable.

Journal Article↗

Intermittency and transient chaos from simple frequency-dependent selection.

Frequency-dependent selection is an important determinant of the evolution of gametophytic self-incompatibility systems in plants, aposematic (warning) and cryptic coloration, systems of mimicry, competitive interactions among members of a population, mating preferences, predator-prey and host-parasite interactions, aggression and other behavioural traits. Past theoretical studies of frequency-dependent selection have shown it to be a plausible mechanism for the maintenance of genetic variability in natural populations. Here, through an analysis of a simple deterministic model for frequency-dependent selection, we demonstrate that complex dynamic behaviour is possible under a broad range of parameter values. In particular we show that the model exhibits not only cycles and chaos but also, for a more restricted set of parameters, transient chaos and intermittency: alterations between an apparently deterministic behaviour and apparently chaotic fluctuations. This behaviour, which has not been stressed within the population genetics literature, provides an explanation for erratic dynamics of gene frequencies.

Alleles↗

Sexual responsiveness is condition-dependent in female guppies, but preference functions are not.

BACKGROUND: Variation in mate choice behaviour among females within a population may influence the strength and form of sexual selection, yet the basis for any such variation is still poorly understood. Condition-dependence may be an important source of variation in female sexual responsiveness and in the preference functions for male display traits that she expresses when choosing. We manipulated food intake of female guppies (Poecilia reticulata), and examined the effect on several measures of condition and various components of mate choice behaviour. RESULTS: Diet significantly influenced four measures of female condition: standard length, weight, reproductive status and somatic fat reserves. Diet also significantly affected female sexual responsiveness, but not preference functions: females in good and poor condition prefer the same males. CONCLUSIONS: Variation in female condition within populations is therefore unlikely to influence the direction of sexual selection imposed by female choice. It may, however, influence the strength of sexual selection due to its effects on female responsiveness. The relative importance of female choice as a sexually selective force may also covary with female condition, however, because low responsiveness may result in sneak copulations being relatively more important as a determinant of the paternity of offspring. Differences among populations in mean condition may also influence geographic differences in the strength of sexual selection.

Analysis of Variance↗

Hormonal status and test condition, but not sexual experience, modulate partner preference in female rats.

A series of experiments was conducted to determine the contributions of hormonal status, test condition, and sexual experience to the display of partner preference by female rats. Preference for a sexually active male rat over a sexually receptive female rat was assessed in independent groups of female rats tested in a condition limiting physical contact (No Contact) and a condition allowing for sexual interaction (Contact). Although hormonal status and test condition influenced the preference for a sexually active male, repeated testing and sexual experience had no effect. Experiment 1 demonstrated that independent of test condition, preference for the male is stronger in estrogen- and progesterone-primed rats than in rats receiving the vehicle. Moreover, independent of hormone condition, rats tested in the No Contact condition exhibit a stronger preference for the male than rats tested in the Contact condition, reflecting in part the active pacing of mating stimulation by sexually receptive rats tested in the Contact condition. Experiment 2 showed that the overall pattern of partner preference in proestrous and diestrous rats was similar to that observed in ovariectomized, estrogen- and progesterone-primed, and oil-treated rats, respectively. In Experiment 3, rats primed with estrogen alone did not exhibit a preference for the male even though fully receptive. Experiments 4 and 5 demonstrated that sexual experience does not affect the expression of preference for the male in estrogen- and progesterone-primed rats. The present findings demonstrate that the female rat's preference for the male is stable across repeated tests and is not affected by sexual experience. Our results also confirm that gonadal hormones influence the expression of a preference for a sexually active male versus a sexually receptive female and demonstrate that the magnitude of preference is modulated by test conditions.

Animals↗

An experimental study of social attraction and spacing between the sexes in sheep.

Most ungulates are gregarious species and outside the mating season are typically observed in single-sex groups. However little is known about the mechanisms underlying social segregation between sexes. We investigated the effect of conspecific attraction on individual spacing between unrestrained merino sheep Ovis aries and confined conspecifics. We considered differences between males and females and whether attractiveness of the confined conspecifics depends on their sex. A series of binary choice experiments was conducted in a large outdoor arena, located in pastures. One or two stimulus animals were placed in small individual cages (1.5 m x 1 m) on opposite sides of the arena. Sheep were tested with one fixed peer of the same or opposite sex vs an empty cage, and with two fixed peers of either the same sex as themselves, or one male and one female. Sheep in a control condition were exposed to two empty cages. In all of the test conditions, confined sheep were highly attractive. Males were more attracted by single stimulus peers of the same than the opposite sex, whereas females did not display such a preference. Sheep confronted with two restrained conspecifics tended to remain between the stimuli. This also occurred when the stimuli were of opposite sex, although the males tended to be located nearer the same-sex peer. Our findings can explain the strong aggregative behaviour of merino sheep, but also the social segregation previously observed in a mixed-sex group through higher attraction for same-sex than opposite-sex peers in males.

Analysis of Variance↗

Maternal naltrexone prevents morphological and behavioral alterations induced in rats by prenatal stress.

The influence of opioid receptor blockade on the developmental and behavioral effects of prenatal stress was studied. Time-mated dams were implanted with minipumps on day 17 of gestation containing vehicle (V) or naltrexone (NTX, 10 mg/kg/day). Noise and light stress was applied on an unpredictable basis, three times a week throughout gestation to half the dams. Maternal NTX completely prevented the reduction in anogenital distance in prenatally stressed (PS) males and restored the growth rate of both sexes. NTX also decreased the anxiety of PS rats in the plus-maze, increased the opioid component of exploration to control levels, but increased anxiety in control males. NTX did not restore the lower saccharin preference in PS females and decreased it in C females. This suggests that some morphological and behavioral changes induced by prenatal stress could result from excess opioid activity induced by maternal stress.

Animals↗

Evolution of sexual isolation during secondary contact: genotypic versus phenotypic changes in laboratory populations.

We monitored the phenotypic and genotypic changes that occur when two behavioral races come into contact in laboratory populations. Drosophila melanogaster from Zimbabwe and nearby regions (Z type) show strong but asymmetric sexual isolation from their cosmopolitan counterparts (M type). Crosses of Z females and M males do not take place readily when other choices are available. At least 15 loci are known to control Z-type mating preferences and performance. By thoroughly mixing the genomes of the two types in laboratory populations, we artificially created maximum secondary contact. Despite the strength of sexual selection favoring Z-type male characters, Z-type behavior is eliminated or greatly diminished in all 12 hybrid populations after only 60 generations. This trend is consistent with the spread of the M-type behavior throughout the world as well as a detailed analysis of fitness components. Surprisingly, in contrast with the phenotypic convergence toward the M-type, genotypic samples broadly covering the genomic regions of mapped behavioral loci show no such trend. The genome appears to be "fine grained," with adjacent loci having different evolutionary dynamics and genealogical histories.

Alleles↗

Interleukin-1 inhibits sexual behavior in female but not in male rats.

The cytokine interleukin-1 (IL-1) is released by a variety of cells in response to infection or injury. IL-1 produces several neuroendocrine and behavioral effects, including a suppression of reproductive functions and goal-directed behaviors. The present study examined the effect of IL-1 on sexual behavior in male and female rats. The following behavioral tests were employed: preference for a sexually appropriate partner, proceptive (soliciting) behavior, the lordosis quotient (sexual receptivity), and mating performance. Peripheral (ip) IL-1 beta, 2 or 10 micrograms/kg, injected 2 h before testing, significantly suppressed proceptive behavior and sexual receptivity in intact, normally cycling females. In ovariectomized rats treated with ovarian hormones, IL-1 beta (2 or 10 micrograms/kg) significantly decreased the preference for a sexually active male partner and suppressed proceptive behavior and sexual receptivity. These effects were evident 2, but not 4 or 6, h after IL-1 beta administration. Intracerebroventricular administration of IL-1 beta (10 ng/rat) also suppressed the preference for a male partner and proceptive behavior in normally cycling females. Similar doses of IL-1 beta had no suppressive effect on any aspect of male sexual behavior, and the highest dose even increased the preference for a receptive female partner. In contrast to the gender-specific effects on sexual behavior, the suppressive effects of IL-1 beta on activity in the open-field test were comparable in male and female rats. The inhibition of female sexual behavior by IL-1 may be adaptive, in that it prevents conception while the animal is sick, thus reducing the risk of spontaneous abortion or abnormal development.

Animals↗

Estimation of assortative mating preferences in the Arctic skua.

Methods are described for the maximum likelihood estimation of mating preferences in models of assortative mating for monogamous and polygamous organisms. These methods are applied to data of matings of the three phenotypes, pale, intermediated and dark of the Arctic Skua. The data were obtained by exhaustive surveys of the Arctic Skua populations on the islands of Fair Isle and Foula. The data give evidence of significant assortative mating of pale birds on Foula and intermediate birds on Fair Isle. The combined data show that there is very highly significant assortative mating, but only of intermediates. In previous surveys, data, in which intermediates and darks were not distinguished, were obtained from a number of islands in the Shetlands. These data, combined with the present data, show that the overall assortative mating of pale is very highly significant with no evidience of heterogeneity. The assortative mating of intermediate birds on Fair Isle agrees with other evidence showing that inermediate males have an advantage as a result of sexual selection.

Animals↗

Behavioral and electrophysiological evidence for volatile sex pheromones in Parcoblatta wood cockroaches.

Species within the cockroach genus Parcoblatta are sexually dimorphic for wing length; females have reduced wings and are flightless, while males have long wings that are used in flight. We predicted that Parcoblatta females would release a volatile sex pheromone to attract the more mobile males. Nymphs of the broad wood cockroach, P. lata, and the Caudell's wood cockroach, P. caudelli, were collected in forested areas in North Carolina, USA, and reared in the laboratory for observations of sexual behavior and for pheromone analysis. After several days of sexual maturation, virgin females of both species exhibited distinct calling behaviors. In females of P. lata, calling commenced 6 days after adult emergence. Under a light-dark photoperiod regime, calling behavior in both species was restricted to the scotophase. Calling consisted of a repeated pattern of raising and lowering the abdomen with occasional exposure of the genital vestibulum. To test whether calling behavior is associated with the release of pheromone, volatiles from calling and noncalling females were collected on Super-Q and tested by electroantennogram (EAG) and behavioral assays. Volatile collections from calling females elicited higher male-specific EAG responses than collections from noncalling females of the same physiological stage. In an olfactometer choice test (Y-tube), males preferred volatiles from calling females over those from noncalling females. To determine the anatomical source of the pheromone, solvent extracts of various body parts were analyzed by EAG. The first through seventh tergites were the only body parts that elicited male-specific EAG responses in both species. In P. lata, the activity of the extract increased from 1- to 7-day-old females, but was lower in mated than in virgin females of the same age. The putative pheromone gland appears to consist of numerous class-3 secretory units, each composed of a secretory cell connected to a cuticular pore via a tubular duct. We conclude that female P. lata and P. caudelli produce ex-specific volatile pheromones that are emitted during calling behavior.

Animal Communication↗

Male mate choice, sexual conflict and strategic allocation of copulations in a lekking bird.

The males of lekking species are not expected to be choosy about mating because a reduced reproductive rate due to lost mating opportunities should outweigh any benefits of male choice. Females have traditionally not been expected to be competitive in this system since their reproduction has usually been assumed to be unconstrained by male availability. Here we show that, in contrast to these predictions, males are choosy and females may be competitive in the lekking great snipe Gallinago media. Males preferred by many females often refused to copulate with and even chased away females that the male had already copulated with, whereas females seemed to compete for repeated copulations. We conclude that choosiness may sometimes pay for popular males in those lekking species where females copulate repeatedly. Apparently, evolutionary conflicts of interest between individuals may cause a richer repertoire of behavioural adaptations than, to our knowledge, hitherto realized.

Animals↗

Morphological adaptation influences the evolution of a mating signal.

Theory predicts that forces of natural selection can reduce the intensity of sexually selected traits. In this study, I investigate how morphological adaptation to feeding ecology influences a mating signal. In birds, changes in feeding ecology can cause rapid divergence in bill morphology. Because bills are also important for song production, feeding ecology may influence song divergence. During song, birds can rapidly change vocal tract resonance using bill movement, yet are constrained in rate and magnitude of bill movements resulting in a trade-off between trill rate and frequency bandwidth. Male swamp sparrows vary in their ability to produce rapid, broad-band trills and females prefer more physically demanding songs. Populations of swamp sparrows adapted to the feeding ecology of tidal marshes have larger bills than inland populations. Larger bills should increase the constraints of producing rapid, broad-band trills allowing for a test of how changes in feeding ecology affect a feature of song used in mate choice. I found significant differences in acoustic features of song consistent with the hypothesis that coastal males are less able to meet the physical demands of song production because of the constraints of having larger bills. As possible compensation for decreases in song performance, coastal populations exhibit an increase in song complexity. These changes support the current model of how motor constraints influence song production and suggest a mechanism by which feeding ecology can influence signal evolution.

Adaptation, Physiological↗