Female Sexual Selection: How Females Choose Mates in the Animal Kingdom

Female sexual selection, also called intersexual selection, is the process by which females of a species choose which males to mate with based on particular traits, behaviors, or signals. This article explains the mechanisms of female choice, presents documented examples across animal groups, and gives students and researchers a practical framework for identifying and studying female choice in wild or captive populations. The content draws on peer-reviewed studies in evolutionary biology and behavioral ecology, with attention to the limits of current evidence and the conditions under which female choice operates.

Defining Female Choice and Its Place in Sexual Selection Theory

Sexual selection operates through two distinct mechanisms that often act on the same traits. Male-male competition involves direct contests between males for access to females, while female mate choice involves females evaluating and selecting among available males. Research on sexual selection has historically focused on one mechanism at a time, but this approach can produce an incomplete picture when the two mechanisms oppose each other or differ in form. Studies that examine both mechanisms together show that male-male competition and female mate choice can reinforce each other or work in opposite directions, and they are more likely to oppose each other when they operate sequentially instead of simultaneously. Body size is the trait most commonly studied through both mechanisms, and selection on body size is typically linear in form. Researchers need standardized measures of the strength and form of selection imposed by each mechanism to understand how they combine into total sexual selection. This matters for practical study design because measuring only one mechanism can lead to incorrect conclusions about how a trait evolves.

Female choice is not a single uniform process. It ranges from simple preferences for larger body size to complex evaluations of multiple signals such as color, song, chemical cues, and behavioral displays. The evolutionary consequences of female choice depend on whether the preferred traits signal genetic quality, whether preferences carry direct costs, and whether the benefits of choice outweigh those costs.

The Good Genes Hypothesis

The good genes hypothesis proposes that females choose males whose traits indicate superior genetic quality, and that offspring of choosy females inherit those benefits. Under this model, exaggerated male ornaments act as honest signals of heritable viability differences. When females choose mates using traits that correlate with heritable viability, stable exaggeration of both female choice and the preferred male trait is possible, even when choice itself is costly. Offspring of choosy females gain both a reproductive advantage through inherited attractiveness and greater viability through inherited quality.

The good genes model faces a theoretical challenge. Female preferences might deplete genetic variance in fitness, making further choice redundant. Additionally, high-condition males sometimes produce low-fitness offspring. This can happen because of environmental turnover and gene-by-environment interactions, where a genotype that performs well in one environment performs poorly in another. It can also happen when fit males carry sexually antagonistic alleles that produce unfit daughters. Evolutionary simulations that incorporate both gene-by-environment interactions and intralocus sexual conflict show that conflict between the sexes can weaken female preferences for high-condition males or even cause preferences for low-condition males. The direction of preference depends on the relative benefits of producing well-adapted sons versus well-adapted daughters, which in turn depends on how strongly selection acts on each sex. This means that the good genes hypothesis cannot be assumed to operate in every species or every environment.

The Runaway Process

The runaway process, originally formalized by Ronald Fisher, describes how female preferences and male traits can coevolve in a self-reinforcing cycle. In Fisher's model, female mating preferences are not subject to direct selection but evolve because they are genetically correlated with the favored male trait. Females who prefer males with exaggerated traits produce sons who inherit those traits and are therefore attractive to the next generation of females. This creates a feedback loop where both the male trait and the female preference become more extreme over time.

The runaway process has an important limitation. When female choice is costly relative to random mating, in terms of energy, time, or predation risk, the evolution of female preference is subject to direct selection. With costly female choice, the line of equilibria found in Fisher's original model no longer exists. Any cost to choice causes choosiness to decline, which lowers the strength of sexual selection and causes the male trait to decline as well. When Fisher's process is the sole force of sexual selection and female choice is costly, only transitory increases in female choice and the preferred male trait are possible. This is why the good genes mechanism is often invoked to explain stable exaggeration of male ornaments in species where female choice carries significant costs.

Phenotypic models of sexual selection offer an alternative way to characterize the evolutionary endpoints of the coevolutionary process. In these models, the mate-choice strategy of female population members determines how attractive females should find each male, and a population is evolutionarily stable if population members actually behave in this way. This approach has practical advantages for computing evolutionarily stable mate-choice strategies, especially when strategies are complex time-dependent preference rules. The sexy son phenomenon, where females prefer males because their sons will inherit attractiveness, can occur even when the male trait is inherited phenotypically instead of genetically, such as through cultural transmission.

At a Glance: Documented Examples of Female Choice Across Taxa

Chemical Signals and Female Choice

Chemical communication plays a central role in female mate choice across the animal kingdom. Pheromones are chemical signals used by members of the same species, and they operate in a wide range of biological contexts, from trail, alarm, and queen pheromones in social insects to the mammary pheromone produced by mother rabbits. Charles Darwin himself proposed that the breeding season sexual smells of male crocodiles, goats, and other animals could have evolved by sexual selection of the smelliest males through female choice. In model organisms such as moths, fruit flies, roundworms, and house mice, complete signaling systems can be genetically dissected, from the enzymes that produce pheromones, through perception by chemosensory receptors, to the neural circuits that process the signals.

The most direct evidence for chemical influence on human mate choice comes from a study of MHC-associated body odors. The major histocompatibility complex is a set of genes involved in immune function, and one proposed benefit of sexual reproduction is that it allows animals to react rapidly to continuously changing environmental selection pressures such as coevolving parasites. This counteraction would be most efficient if females could provide their progeny with certain allele combinations for loci that may be crucial in the parasite-host arms race, such as the MHC. In a study of university students, male students wore T-shirts for two consecutive nights, and female students rated the odors of six T-shirts each. Women scored male body odors as more pleasant when the men differed from them in MHC type than when they were more similar. This difference in odor assessment was reversed when the women rating the odors were taking oral contraceptives. Furthermore, the odors of MHC-dissimilar men reminded the test women more often of their own actual or former mates than did the odors of MHC-similar men. This suggests that the MHC or linked genes influence human mate choice today.

Visual Signals and Female Choice in Fish

Fish provide some of the most tractable systems for studying female choice because their visual signals can be manipulated experimentally and their preferences can be measured in controlled laboratory settings.

Guppies and Mate-Choice Copying

Female guppies engage in mate-choice copying, where the presence and number of other females choosing a particular male influences a female's own preference. A 2025 study in the Journal of Ethology examined how the number of models affects mate-choice copying in female guppies. The title of the study, "Number matters: effect of the number of models on mate-choice copying in female guppies," indicates that the quantity of social information available affects the strength of copying behavior. This social learning mechanism allows females to reduce the costs of individual assessment by relying on the choices of other females who may have better information about male quality.

Sticklebacks and Video Playback

Video playback experiments have been used to study female mate choice in sticklebacks, with a 1995 study examining female motivation and attentiveness to male color cues. The use of video playback allows researchers to isolate specific visual traits and present them to females in a controlled manner, eliminating confounding variables such as male behavior or chemical signals. This method has proven valuable for determining which specific color cues females attend to and how motivated they are to approach males displaying those cues.

Western Mosquitofish and 3D Simulation

A 2024 study used 3D simulation technology to investigate mate choice in western mosquitofish. Researchers created 3D simulation animations using Maya 2018 software to accurately replicate key movements and behaviors essential for the research. Through preference tests, they validated these animations and found that the fish could effectively identify 3D simulated mates. Both male and female mosquitofish exhibited a strong preference for larger animations. The fish displayed significantly greater attraction to 3D simulations compared to 2D ones. This work demonstrates the potential of 3D simulation technology for studying fish behavior, offering an efficient, precise, and non-invasive method for future research on mate choice.

Green Terror Cichlids and Sex Differences in Preference

A 2022 study of the green terror cichlid (Aequidens rivulatus) analyzed differences in mate choice behavior and criteria between males and females. The study quantified body size, behavioral intention, and appearance through a no-choice paradigm. Results showed that males paid more attention to preference degree and female attractiveness, whereas females focused on ability and physical strength displays. Males who chose to mate were primarily associated with body size, behavioral intention, and appearance, whereas the preferences of females were body size, appearance, and behavioral intention. These findings reveal that males and females have different criteria for mate choice, which is vital for determining successful mating and improving artificial mating in aquaculture settings.

Acoustic Signals and Female Choice in Frogs

Acoustic communication is a primary channel for female mate choice in many frog species. Female serrate-legged small treefrogs (Kurixalus odontotarsus) were tested for mate choice decisions under different ambient light conditions. Artificial light at night is a widespread anthropogenic stimulus that can significantly alter nocturnal animal behavior, from migration to foraging to vocal communication. Researchers tested the hypothesis that mate choice decisions of female serrate-legged small treefrogs were influenced by ambient light intensity using standard two-speaker phonotaxis tests in a sound attenuating chamber. They set four light conditions ranging from the maximum natural light at night, equivalent to a full moon, to that of actual calling sites with artificial light.

Contrary to prediction, female frogs showed a preference for calls on the bright side when exposed to identical stimuli under the lowest light condition. However, females preferred longer calls on the dim side to shorter calls on the bright side in this treatment. There were no significant effects of choice side, light treatment, or their interaction on leave time or choice time. These results suggest that females are more attracted to mates in bright light under natural nocturnal light conditions, but the preference for longer calls is not altered in serrate-legged small treefrogs. This study demonstrates that environmental conditions can influence mate choice behavior without necessarily altering the underlying preference for acoustic traits.

Social Context and the Interaction of Competition and Choice

Female choice does not operate in isolation. The social environment, including the presence and behavior of other individuals, can dramatically alter how female preferences are expressed.

Maritime Earwigs and Group Composition

A 2026 study of the maritime earwig (Anisolabis maritima) examined how sex and body size influence aggression and courtship in different-sized groups. This insect species has males and females that differ morphologically and behaviorally. Researchers videotaped mixed-sex pairs and trios of earwigs for two hours to determine the effects of sex and size on their interactions. In pairs, females were more aggressive than males, and both sexes showed size-based aggression. However, sex was a stronger determinant of aggression in male-biased trios, whereas sex and size were more important in female-biased trios.

Male size did not affect courtship in pairs, as large and small males were equally likely to engage in copulatory activity. The presence of intrasexual selection altered the results in the two types of trios. In male-biased trios, the larger male had more copulatory opportunities as his size relative to his rival increased, indicating that intrasexual domination by larger males can overcome female preferences for smaller males observed in previous studies. In female-biased trios, there were no such patterns since copulatory opportunities were relatively rare, the likely result of elevated aggression with two interacting females. These results indicate that sex and size affect both aggression and sexual behavior differently based on group size and composition, which highlights the need to examine behaviors in a variety of social contexts.

Medaka and Oxytocin Regulation

A 2026 study in medaka fish investigated how oxytocin regulates familiarity-dependent mate choice. Some social animals have evolved the ability to distinguish between familiar and unfamiliar individuals and to flexibly modify their behavior according to social context. Disruption of this system has been implicated in neurodevelopmental disorders such as autism, motivating efforts to elucidate its molecular basis. In female medaka, wild-type individuals preferentially accepted familiar males, whereas oxytocin-deficient females rapidly accepted even unfamiliar males, resulting in a loss of familiarity-dependent preference. In contrast, oxytocin-deficient males showed reduced courtship toward unfamiliar females, but their courtship frequency increased with cohabitation. In triadic social interactions, oxytocin-deficient males displayed enhanced mate-guarding behavior toward familiar females, indicating exaggerated preference.

These results demonstrate that oxytocin deficiency produces sex-dependent effects on mate choice. Whole-brain transcriptome analyses revealed a consistent reduction in the expression of complement component C1q genes in both sexes, suggesting abnormalities in neurodevelopmental processes. Genes related to GABA metabolism were selectively altered in females, indicating sex-specific molecular pathways downstream of oxytocin signaling. This study illustrates how molecular mechanisms can regulate the social recognition systems that underlie female choice.

Genital Evolution and Cryptic Female Choice

Female choice extends beyond pre-copulatory decisions. Cryptic female choice occurs after mating begins and can involve female-controlled mechanisms that influence which male's sperm fertilizes her eggs. Genital coevolution between the sexes is expected to be common because of the direct interaction between male and female genitalia during copulation. The diverse mechanisms of genital coevolution include natural selection, female mate choice, male-male competition, and their interactions, which can generate sexual conflict that leads to sexually antagonistic coevolution.

Natural selection on genital morphology results in size coevolution to allow copulation to be mechanically possible, even as other features of genitalia may reflect the action of other selection mechanisms. Genital coevolution is explicitly predicted by at least three mechanisms of genital evolution: lock and key to prevent hybridization, female choice, and sexual conflict. Although some good examples exist in support of each mechanism, more data on quantitative female genital variation and studies of functional morphology during copulation are needed to understand more general patterns. A combination of approaches is required to advance understanding of genital coevolution, including knowledge of the ecology and behavior of the studied species combined with functional morphology, quantitative morphological tools, experimental manipulation, and experimental evolution. The best-studied species are all invertebrates, so attention to vertebrates in any of these areas is badly needed.

Female Choice in Primates and Humans

Mandrills as a Model System

Mandrills (Mandrillus sphinx) are a classic example of extravagant armaments and ornaments in animals. Long-term, multidisciplinary research that integrates field observations with laboratory methods has contributed to ongoing theoretical debates in the field of sexual selection. Mandrill life history, the ontogeny of sex differences, and maternal effects all shape the expression of sexually selected traits. Male-male competition and female choice both operate in this species, and less well-studied questions of female-female competition and male choice are also relevant.

Different reproductive priorities lead to very different life histories and divergent adaptations in males and females. Broadening traditional perspectives on sexual selection beyond the ostentatious results of intense sexual selection on males leads to an understanding of more subtle and cryptic forms of competition and choice in both sexes. This opens productive avenues in the study of primate reproductive strategies, including postcopulatory selection, female intrasexual competition, and male choice. Studies of mandrills provide comparison and inspiration for studies of both other polygynandrous species and species with mating systems less traditionally associated with sexual selection.

Human Mate Choice and MHC

The MHC study described earlier provides evidence that human female mate choice is influenced by immune gene compatibility. The finding that oral contraceptive use reverses the preference pattern is particularly important because it suggests that hormonal status affects how women perceive MHC-related odor cues. This has implications for understanding mate choice in modern human populations where hormonal contraception is widely used.

Practical Framework for Studying Female Choice

Researchers and students who want to study female choice in their own study systems should follow a structured approach that accounts for the complexity of the process.

Step 1: Define the Trait and the Preference

Identify the male trait or traits that females might be evaluating. This requires detailed behavioral observation and, where possible, manipulation of the trait. Common traits include body size, coloration, acoustic signals, chemical signals, and behavioral displays. The trait must vary among males for choice to be possible.

Step 2: Design Choice Experiments

Choice experiments typically present a female with two or more males or simulated male stimuli and measure her approach, association time, or other indicators of preference. Methods range from simple two-choice tests to video playback and 3D simulation. Each method has tradeoffs. Live males provide natural behavior but introduce confounding variables. Video playback controls stimuli but may lack natural cues. 3D simulation offers precise control while maintaining realistic movement.

Step 3: Measure Both Mechanisms of Sexual Selection

Do not measure female choice in isolation. Quantify male-male competition acting on the same trait to determine whether the two mechanisms reinforce or oppose each other. This requires observing interactions among males as well as female responses. The timing of the two mechanisms matters, since they are more likely to oppose each other when they operate sequentially.

Step 4: Assess the Costs of Choice

Determine whether female choice carries costs in energy, time, or predation risk. If choice is costly, the good genes mechanism is more likely to maintain stable preferences. If choice is costless, Fisherian runaway can operate. Measuring costs requires comparing the behavior of choosy females with females forced to mate randomly.

Step 5: Test for Genetic Benefits

To distinguish between good genes and Fisherian processes, measure the fitness of offspring produced by choosy versus non-choosy females. This requires breeding designs that control for maternal effects and environmental variation. Gene-by-environment interactions can complicate these measurements, so offspring fitness should be assessed across relevant environmental conditions.

Step 6: Consider Social Context

Female preferences expressed in one social context may not hold in another. The earwig study demonstrates that the presence of competing males or other females can override or alter female preferences. Study female choice across multiple group sizes and compositions before drawing conclusions about the species.

Records and Measurements

Maintain standardized records for female choice studies. For each trial, record the female identity, the stimuli presented, the duration of the trial, the response variable measured, and the environmental conditions. For live male stimuli, record male identity, trait values, and behavioral state. For simulated stimuli, record the specific parameters of the simulation.

Quantitative measures of female choice include association time, number of approaches, latency to approach, and phonotaxis responses in acoustic studies. For studies of mate-choice copying, record the number and identity of model females and the order of choices. For studies of social context, record group composition and the timing of interactions.

Common Failure Patterns in Female Choice Research

Several recurring problems compromise female choice studies. The most common is measuring only one mechanism of sexual selection and attributing all selection to that mechanism. This produces misleading conclusions when the mechanisms oppose each other. A second failure is ignoring the costs of choice, which leads to incorrect predictions about evolutionary stability. A third failure is studying female choice in a single social context and generalizing to all contexts. A fourth failure is neglecting gene-by-environment interactions, which can reverse the direction of preference under different environmental conditions. A fifth failure is using simulated stimuli without validating that females respond to them as they respond to live males.

Limitations of Current Evidence

The evidence base for female choice is uneven across taxa. Invertebrates, particularly insects, are overrepresented in studies of genital coevolution and chemical communication. Vertebrates, especially mammals and birds, are underrepresented in experimental studies of genital evolution. Long-term studies that integrate field observations with laboratory methods, such as the mandrill research, are rare but provide the most complete picture. Standardized measures of the form and strength of selection imposed by each mechanism of sexual selection are lacking across most species.

The theoretical literature identifies conditions under which female choice should or should not evolve, but empirical tests of these predictions are incomplete. The interaction between gene-by-environment interactions and intralocus sexual conflict in shaping female preferences has been modeled but not thoroughly tested empirically. The role of social learning in female choice, including mate-choice copying, is documented in some species but the mechanisms and limits of this learning are not fully understood.

Welfare and Ethical Considerations

Studies of female choice involve live animals and require attention to welfare. Choice experiments should minimize stress and avoid prolonged confinement. Simulated stimuli, including video playback and 3D animation, offer non-invasive alternatives that reduce the need for live animal interactions. When live animals are used, housing conditions should match species-appropriate requirements, and trials should be terminated if animals show signs of distress.

For species used in aquaculture or captive breeding, understanding female choice has practical applications. The green terror cichlid study explicitly notes that understanding mate choice criteria is vital for improving artificial mating. Breeders can use knowledge of female preferences to design pairing strategies that improve reproductive success and maintain genetic diversity.

Professional Escalation Criteria

Researchers studying female choice should seek consultation with statisticians or evolutionary theorists when their data show unexpected patterns, such as preferences that reverse direction across contexts or environments. Studies involving molecular techniques, such as the oxytocin work in medaka, require specialized expertise and should be conducted in collaboration with molecular biologists. Studies of human mate choice involving hormonal status require ethical approval and should be designed with attention to participant welfare and confidentiality.

When results contradict established theory, such as finding stable preferences in the absence of good genes benefits, researchers should consider whether unmeasured costs or benefits are operating. When results contradict previous findings in the same species, such as the earwig study where intrasexual competition overrode female preferences, researchers should examine whether social context explains the discrepancy.

Frequently Asked Questions

What is the difference between female choice and male-male competition?

Female choice, also called intersexual selection, is the process where females evaluate and select mates based on particular traits. Male-male competition, also called intrasexual selection, is the process where males compete directly with each other for access to females. Both mechanisms can act on the same trait, and they can reinforce or oppose each other. Studies that measure only one mechanism can produce incomplete or misleading conclusions about how sexually selected traits evolve.

How do researchers distinguish between good genes and runaway selection?

Good genes selection is indicated when preferred male traits correlate with heritable viability differences and offspring of choosy females show greater survival or fitness. Runaway selection is indicated when female preferences evolve because they are genetically correlated with the favored male trait, regardless of whether the trait signals genetic quality. The two processes can be distinguished by measuring offspring fitness and by testing whether preferences persist when choice is costly. When female choice is costly, stable exaggeration of male traits is more likely to require good genes benefits.

Why do females sometimes prefer males that do not provide direct benefits?

Females may prefer males that provide indirect genetic benefits, such as genes for parasite resistance or attractiveness that are passed to offspring. The good genes hypothesis proposes that preferred traits signal genetic quality. The runaway process proposes that preferences evolve because of genetic correlation with the male trait, producing sexy sons who are attractive to the next generation of females. In some cases, preferences may be maintained by social learning, such as mate-choice copying, where females use the choices of other females as information.

How does social context affect female choice?

Social context can dramatically alter female choice. The maritime earwig study showed that in male-biased groups, larger males dominated copulatory opportunities and overcame female preferences for smaller males. In female-biased groups, aggression between females reduced copulatory opportunities overall. Group size and composition affect both aggression and sexual behavior differently, so female preferences observed in one social context may not hold in another.

What role do chemical signals play in female choice?

Chemical signals, including pheromones, are used by species across the animal kingdom in a wide range of biological contexts. In humans, MHC-related body odors influence female preferences, with women rating odors of MHC-dissimilar men as more pleasant. This preference is reversed when women take oral contraceptives. In model organisms such as moths, fruit flies, roundworms, and house mice, complete pheromone signaling systems can be genetically dissected from production to perception to neural processing.

Can female choice be studied using simulated stimuli?

Yes, simulated stimuli are increasingly used in female choice research. Video playback experiments have been used to study stickleback mate choice, and 3D simulation technology has been validated for studying western mosquitofish. Both male and female mosquitofish exhibited strong preferences for larger animations and showed greater attraction to 3D simulations compared to 2D ones. Simulated stimuli offer precise control over traits and reduce the need for live animal interactions, but they must be validated to ensure females respond to them as they respond to live males.

How does hormonal status affect female mate choice?

Hormonal status can influence female preferences. In the human MHC study, women's preferences for MHC-dissimilar body odors were reversed when the women were taking oral contraceptives. In medaka fish, oxytocin deficiency eliminated the normal female preference for familiar males, while producing different effects in males. These findings indicate that hormonal and molecular pathways regulate the expression of female preferences and that these pathways can differ between the sexes.

Why is it important to study both sexes in mate choice research?

Males and females often have different mate choice criteria. The green terror cichlid study found that males paid more attention to preference degree and female attractiveness, whereas females focused on ability and physical strength displays. Different reproductive priorities lead to divergent adaptations in males and females. Studying both sexes reveals more subtle and cryptic forms of competition and choice and provides a more complete understanding of reproductive strategies.

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References and Further Reading

• NCBI Literature Resources. National Center for Biotechnology Information.
• PubMed. National Library of Medicine.
• MHC-dependent mate preferences in humans.. Proceedings. Biological sciences, 1995.
• Pheromones.. Current biology : CB, 2017.
• Male-male competition, female mate choice and their interaction: determining total sexual selection.. Journal of evolutionary biology, 2009.
• Evolution of female choice under intralocus sexual conflict and genotype-by-environment interactions.. Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 2018.
• The costs of choice in sexual selection.. Journal of theoretical biology, 1987.
• Sexual Selection and the differences between the sexes in Mandrills (Mandrillus sphinx).. American journal of physical anthropology, 2016.
• Quantifying male attractiveness.. Proceedings. Biological sciences, 2003.
• Mechanisms and Evidence of Genital Coevolution: The Roles of Natural Selection, Mate Choice, and Sexual Conflict.. Cold Spring Harbor perspectives in biology, 2015.
• Two's company, three's a crowd: Social situations alter group dynamics in the maritime earwig (Anisolabis maritima).. 2026.
• Number matters: effect of the number of models on mate-choice copying in female guppies. Journal of ethology, 2025.
• Video playback experiments on stickleback mate choice: female motivation and attentiveness to male colour cues. 1995.
• Sex Differences in Mate Choice Preference Characteristics of Aequidens rivulatus. Animals, 2022.
• Mate Choice in Western Mosquitofish (Gambusia affinis) in Response to Virtual Mates: A Method for the Investigation of Fish Mate Choice Using Maya 3D Simulation Technology. Animals, 2024.
• Mate choice decisions of female serrate-legged small treefrogs are affected by ambient light under natural, but not enhanced artificial nocturnal light conditions.. Behavioural Processes, 2019.
• [Oxytocin regulates social familiarity-dependent mate choice in medaka].. Nihon yakurigaku zasshi. Folia pharmacologica Japonica, 2026.
• Sexual selection in neotropical spiders: Examples from selected groups. Behaviour and Ecology of Spiders Contributions from the Neotropical Region, 2017.
• Sexual selection by male choice in monogamous and polygynous human populations. Theoretical Population Biology, 1999.
• Male sexual signals and female choice in Drosophila grimshawi (Diptera: Drosophilidae). Journal of Insect Behavior, 1998.

This article is educational and does not replace institutional policy, professional advice, or applicable safety and regulatory requirements.