In the evolutionary biology of sexual reproduction, operational sex ratio (OSR) is the ratio of sexually competing males that are ready to mate to sexually competing females that are ready to mate, [1] [2] [3] or alternatively the local ratio of fertilizable females to sexually active males at any given time. [4] This differs from physical sex ratio which simply includes all individuals, including those that are sexually inactive or do not compete for mates.
The theory of OSR hypothesizes that the operational sex ratio affects the mating competition of males and females in a population. [5] This concept is especially useful in the study of sexual selection since it is a measure of how intense sexual competition is in a species, and also in the study of the relationship of sexual selection to sexual dimorphism. [6] The OSR is closely linked to the "potential rate of reproduction" of the two sexes; [1] that is, how fast they each could reproduce in ideal circumstances. Usually variation in potential reproductive rates creates bias in the OSR and this in turn will affect the strength of selection. [7] The OSR is said to be biased toward a particular sex when sexually ready members of that sex are more abundant. For example, a male-biased OSR means that there are more sexually competing males than sexually competing females.
The operational sex ratio is affected by the length of time each sex spends in caring for young or in recovering from mating. [8] For example, if females cease mating activity to care for young, but males do not, then more males would be ready to mate, thus creating a male biased OSR. One aspect of gestation and recovery time would be clutch loss. Clutch loss is when offspring or a group of offspring is lost, due to an accident, predation, etc. This, in turn, affects how long reproductive cycles will be in both males and females. If the males were to invest more time in the care of their offspring, they would be spending less time mating. This pushes the population towards a female biased OSR and vice versa. Whether or not it is the males or females investing more care in their offspring, if they were to lose their offspring for whatever reason, this would then change the OSR to be less biased because the once occupied sex becomes available to mate again. [9]
As aforementioned, another major factor that influences OSR is potential rate of reproduction (PRR). Any sexual differences in the PRR will also change the OSR, so it is important to look at factors that change PRR as well. [10] [11] [12] [13] These include constraints to environmental factors such as food or nesting sites. For example, if males are required to provide a nutrient high gift before mating (most likely food) then when nutrients available is high, the OSR will be male biased because there is plenty of nutrients available to provide gifts. However, if nutrients is low, less males will be ready to reproduce, causing the population to have a female biased OSR. [10] [14] [15] [16] Another example would be if, in a certain species, males provided care for offspring and a nest. [17] If the availability of nesting sites decreased, we would see the population trend towards a more female biased OSR because only a small number of males actually have a nest while all the females, regardless of a nest or not, are still producing eggs. [18]
A major factor that OSR can predict is the opportunity for sexual selection. As the OSR becomes more biased, the sex that is in excess will tend to undergo more competition for mates and therefore undergo strong sexual selection. [4] [8] [19] Intensity of competition is also a factor that can be predicted by OSR. [2] According to sexual selection theory, whichever sex is more abundant is expected to compete more strongly and the sex that is less abundant is expected to be "choosier" in who they decide to mate with. It would be expected that when an OSR is more biased to one sex than the other, that one would observe more interaction and competition from the sex that is more available to mate. When the population is more female biased, more female-female competition is observed and the opposite is seen for a male population where a male biased would cause more male-male interaction and competitiveness. Though both sexes may be competing for mates, it is important to remember that the biased OSR predicts which sex is the predominant competitor (the sex that exhibits the most competition). [10] [20] [21] OSR can also predict what will happen to mate guarding in a population. As OSR becomes more biased to one sex, it can be observed that mate-guarding will increase. This is likely due to the fact that rival numbers (number of a certain sex that are also ready to mate) are increased. If a population is male biased then there are a lot more rival males to compete for a mate, meaning that those who have a mate already are more likely to guard the mate that they have. [22]
Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with, and compete with members of the same sex for access to members of the opposite sex. These two forms of selection mean that some individuals have greater reproductive success than others within a population, for example because they are more attractive or prefer more attractive partners to produce offspring. Successful males benefit from frequent mating and monopolizing access to one or more fertile females. Females can maximise the return on the energy they invest in reproduction by selecting and mating with the best males.
Sexual dimorphism is the condition where sexes of the same species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most dioecious species, which consist of most animals and some plants. Differences may include secondary sex characteristics, size, weight, color, markings, or behavioral or cognitive traits. Male-male reproductive competition has evolved a diverse array of sexually dimorphic traits. Aggressive utility traits such as "battle" teeth and blunt heads reinforced as battering rams are used as weapons in aggressive interactions between rivals. Passive displays such as ornamental feathering or song-calling have also evolved mainly through sexual selection. These differences may be subtle or exaggerated and may be subjected to sexual selection and natural selection. The opposite of dimorphism is monomorphism, when both biological sexes are phenotypically indistinguishable from each other.
A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. A lek can also indicate an available plot of space able to be utilized by displaying males to defend their own share of territory for the breeding season. A lekking species is characterised by male displays, strong female mate choice, and the conferring of indirect benefits to males and reduced costs to females. Although most prevalent among birds such as black grouse, lekking is also found in a wide range of vertebrates including some bony fish, amphibians, reptiles, and mammals, and arthropods including crustaceans and insects.
Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for animal behavior due to ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined four questions to address when studying animal behaviors: What are the proximate causes, ontogeny, survival value, and phylogeny of a behavior?
A sex ratio is the ratio of males to females in a population. As explained by Fisher's principle, for evolutionary reasons this is typically about 1:1 in species which reproduce sexually. However, many species deviate from an even sex ratio, either periodically or permanently. Examples include parthenogenic species, periodically mating organisms such as aphids, some eusocial wasps, bees, ants, and termites.
Sperm competition is the competitive process between spermatozoa of two or more different males to fertilize the same egg during sexual reproduction. Competition can occur when females have multiple potential mating partners. Greater choice and variety of mates increases a female's chance to produce more viable offspring. However, multiple mates for a female means each individual male has decreased chances of producing offspring. Sperm competition is an evolutionary pressure on males, and has led to the development of adaptations to increase male's chance of reproductive success. Sperm competition results in a sexual conflict between males and females. Males have evolved several defensive tactics including: mate-guarding, mating plugs, and releasing toxic seminal substances to reduce female re-mating tendencies to cope with sperm competition. Offensive tactics of sperm competition involve direct interference by one male on the reproductive success of another male, for instance by physically removing another male's sperm prior to mating with a female. For an example, see Gryllus bimaculatus.
In evolutionary biology and evolutionary psychology, the Trivers–Willard hypothesis, formally proposed by Robert Trivers and Dan Willard in 1973, suggests that female mammals adjust the sex ratio of offspring in response to maternal condition, so as to maximize their reproductive success (fitness). For example, it may predict greater parental investment in males by parents in "good conditions" and greater investment in females by parents in "poor conditions". The reasoning for this prediction is as follows: Assume that parents have information on the sex of their offspring and can influence their survival differentially. While selection pressures exist to maintain a 1:1 sex ratio, evolution will favor local deviations from this if one sex has a likely greater reproductive payoff than is usual.
Sex allocation is the allocation of resources to male versus female reproduction in sexual species. Sex allocation theory tries to explain why many species produce equal number of males and females.
Sexual conflict or sexual antagonism occurs when the two sexes have conflicting optimal fitness strategies concerning reproduction, particularly over the mode and frequency of mating, potentially leading to an evolutionary arms race between males and females. In one example, males may benefit from multiple matings, while multiple matings may harm or endanger females, due to the anatomical differences of that species. Sexual conflict underlies the evolutionary distinction between male and female.
Mate choice is one of the primary mechanisms under which evolution can occur. It is characterized by a "selective response by animals to particular stimuli" which can be observed as behavior. In other words, before an animal engages with a potential mate, they first evaluate various aspects of that mate which are indicative of quality—such as the resources or phenotypes they have—and evaluate whether or not those particular trait(s) are somehow beneficial to them. The evaluation will then incur a response of some sort.
The sexy son hypothesis in evolutionary biology and sexual selection, proposed by Patrick J. Weatherhead and Raleigh J. Robertson of Queen's University in Kingston, Ontario in 1979, states that a female's ideal mate choice among potential mates is one whose genes will produce males with the best chance of reproductive success. This implies that other benefits the father can offer the mother or offspring are less relevant than they may appear, including his capacity as a parental caregiver, territory and any nuptial gifts. Fisher's principle means that the sex ratio is always near 1:1 between males and females, yet what matters most are her "sexy sons'" future breeding successes, more likely if they have a promiscuous father, in creating large numbers of offspring carrying copies of her genes. This sexual selection hypothesis has been researched in species such as the European pied flycatcher.
A courtship display is a set of display behaviors in which an animal, usually a male, attempts to attract a mate; the mate exercises choice, so sexual selection acts on the display. These behaviors often include ritualized movement ("dances"), vocalizations, mechanical sound production, or displays of beauty, strength, or agonistic ability.
Male-male intrasexual competition occurs when two males of the same species compete for the opportunity to mate with a female. Sexually dimorphic traits, size, sex ratio, and the social situation may all play a role in the effects male-male competition has on the reproductive success of a male and the mate choice of a female. Larger males tend to win male-male conflicts due to their sheer strength and ability to ward off other males from taking over their females. For instance, in the fly Dryomyza anilis, size shows the strongest correlation to the outcome of male-male conflicts over resources like territory and females.
Monogyny is a specialised mating system in which a male can only mate with one female throughout his lifetime but the female may mate with more than one male. In this system the males generally provide no paternal care. In many spider species that are monogynous, the males have two copulatory organs, which allows them to mate a maximum of twice throughout their lifetime. As is commonly seen in honeybees, ants and certain spider species, a male may put all his energy into a single copulation, knowing that this will lower his overall fitness. During copulation monogynous males have adapted to cause self genital damage or even death to increase their chances of paternity.
Bateman's principle, in evolutionary biology, is that in most species, variability in reproductive success is greater in males than in females. It was first proposed by Angus John Bateman (1919–1996), an English geneticist. Bateman suggested that, since males are capable of producing millions of sperm cells with little effort, while females invest much higher levels of energy in order to nurture a relatively small number of eggs, the female plays a significantly larger role in their offspring's reproductive success. Bateman's paradigm thus views females as the limiting factor of parental investment, over which males will compete in order to copulate successfully.
Sexual selection in birds concerns how birds have evolved a variety of mating behaviors, with the peacock tail being perhaps the most famous example of sexual selection and the Fisherian runaway. Commonly occurring sexual dimorphisms such as size and color differences are energetically costly attributes that signal competitive breeding situations. Many types of avian sexual selection have been identified; intersexual selection, also known as female choice; and intrasexual competition, where individuals of the more abundant sex compete with each other for the privilege to mate. Sexually selected traits often evolve to become more pronounced in competitive breeding situations until the trait begins to limit the individual's fitness. Conflicts between an individual fitness and signaling adaptations ensure that sexually selected ornaments such as plumage coloration and courtship behavior are "honest" traits. Signals must be costly to ensure that only good-quality individuals can present these exaggerated sexual ornaments and behaviors.
Sexual selection in mammals is a process the study of which started with Charles Darwin's observations concerning sexual selection, including sexual selection in humans, and in other mammals, consisting of male–male competition and mate choice that mold the development of future phenotypes in a population for a given species.
Sexual selection in scaled reptiles studies how sexual selection manifests in snakes and lizards, which constitute the order Squamata of reptiles. Each of the over three thousand snakes use different tactics in acquiring mates. Ritual combat between males for the females they want to mate with includes topping, a behavior exhibited by most viperids in which one male will twist around the vertically elevated fore body of its opponent and forcing it downward. It is common for neck biting to occur while the snakes are entwined.
Formica truncorum is a species of wood ant from the genus Formica. It is distributed across a variety of locations worldwide, including central Europe and Japan. Workers can range from 3.5 to 9.0mm and are uniquely characterized by small hairs covering their entire bodies. Like all other ants, F. truncorum is eusocial and demonstrates many cooperative behaviors that are unique to its order. Colonies are either monogynous, with one queen, or polygynous, with many queens, and these two types of colonies differ in many characteristics.
Inbreeding avoidance, or the inbreeding avoidance hypothesis, is a concept in evolutionary biology that refers to the prevention of the deleterious effects of inbreeding. Animals only rarely exhibit inbreeding avoidance. The inbreeding avoidance hypothesis posits that certain mechanisms develop within a species, or within a given population of a species, as a result of assortative mating and natural and sexual selection, in order to prevent breeding among related individuals. Although inbreeding may impose certain evolutionary costs, inbreeding avoidance, which limits the number of potential mates for a given individual, can inflict opportunity costs. Therefore, a balance exists between inbreeding and inbreeding avoidance. This balance determines whether inbreeding mechanisms develop and the specific nature of such mechanisms.