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Polygyny threshold model

The polygyny threshold model is an evolutionary framework in behavioral ecology that predicts females will tolerate polygyny—mating with a male already paired with other females—only if the net fitness gains from his superior resources or territory exceed the fitness achievable through monogamy with an unpaired, lower-quality male, thereby defining a resource-based "threshold" for adaptive polygynous choice.^[1] Formulated by Gordon H. Orians in 1969, the model assumes greater female parental investment leads to choosiness, with polygyny arising when ecological variability in male quality creates disparities where sharing a top-tier male outperforms solo pairing with inferiors.^[1]^[2] The model's core graphical representation plots female fitness against male resource quality, showing a monogamy curve for primary females and a lower, parallel polygyny curve for secondaries, with the threshold as the point of intersection where choices equilibrate.^[1] Empirical validation includes experimental manipulations in red-winged blackbirds (Agelaius phoeniceus), where females preferred high-quality territories despite existing mates, confirming compensatory benefits offset sharing costs like reduced paternal care.^[3] Extensions to mammals highlight similar dynamics, with polygyny favored in resource-heterogeneous environments.^[2] In human societies, the model illuminates historical polygyny prevalence—documented in over 80% of preindustrial cultures—where wealthy males monopolize mates, potentially elevating female reproductive success via resource access, though generalized versions incorporating male choosiness forecast reduced polygyny under extreme wealth inequality due to surplus low-status males competing unsuccessfully.^[4]^[5] Defining characteristics include its emphasis on female agency in mating decisions driven by fitness maximization, rather than male coercion, challenging assumptions of inherent female detriment in polygyny.^[6] Notable controversies encompass reassessments questioning static thresholds amid temporal habitat variability or offspring quality trade-offs, and critiques positing overlooked sexual conflicts where male interests diverge from female optima.^[7]^[8] Despite modifications, the model remains foundational for understanding mating system evolution, privileging causal links between ecology, sex roles, and reproductive strategies over normative cultural interpretations.^[9]

Origins and Historical Development

Initial Proposal by Gordon Orians

Gordon H. Orians introduced the polygyny threshold model in his 1969 paper "On the Evolution of Mating Systems in Birds and Mammals," framing it within a theory of female mate selection aimed at maximizing individual reproductive success.^[1] The model addresses resource-defense polygyny, where males control territories varying in quality due to habitat heterogeneity, and females must decide between monogamous pairing with a lower-quality male on a suboptimal territory or sharing a higher-quality male and territory. Orians posited that females should enter polygynous unions only if the fitness benefits from superior resources exceed the costs of divided male parental care and resource allocation, defining a "threshold" as the critical disparity in territory productivity that tips the balance toward polygyny.^[10] Central assumptions include greater female parental investment, making females the sex-limited resource; male defense of territories as proxies for female reproductive potential; and a inherent cost to polygyny for females on equivalent territories, stemming from reduced per-female resource access and male assistance. Orians emphasized ecological drivers, predicting polygyny's prevalence in contexts of pronounced resource variation, such as mammals (where male investment is lower) over birds, precocial over altricial birds, and habitats like marshes or early-successional stages with patchy, defensible resources over uniform terrestrial ones.^[1] The model also forecasts polyandry's rarity, as role reversals in investment are uncommon, and links polygyny to traits like widespread foraging grounds paired with restricted nesting sites.^[1] This economic approach treated female choice as adaptive optimization, integrating habitat quality with male attributes to explain polygyny's persistence despite apparent monogamy biases in many taxa.^[1] Orians' predictions aligned partially with observed patterns, such as higher polygyny rates in mammals (around 10-15% of species versus under 5% in birds), though incomplete fits highlighted needs for refinement, including imperfect information and genetic factors.^[1] The proposal shifted focus from male-male competition alone to female-driven evolutionary dynamics, influencing behavioral ecology by underscoring causal links between environmental variance and mating strategies.^[2] In 1977, Stephen T. Emlen and Lewis W. Oring integrated the polygyny threshold model into a broader ecological theory of mating system evolution, positing that mating strategies arise from constraints on access to critical resources and mates, with females evaluating the net fitness benefits of polygyny based on male-controlled breeding conditions versus unpaired alternatives.^[11] Their framework emphasized the operational sex ratio and differential parental investment as factors modulating the threshold, predicting polygyny where high-quality males monopolize superior habitats, forcing some females to share mates rather than settle for inferior monogamous options.^[11] This refinement shifted focus from static territory quality to dynamic ecological pressures, including temporal variability in resource availability that influences female choice timing.^[11] James F. Wittenberger further developed the model in the late 1970s and early 1980s by incorporating temporal dimensions, such as the sequence of male territory establishment and female mate searching, which he argued could alter the effective polygyny threshold by affecting female knowledge of habitat options.^[12] In a 1981 analysis, Wittenberger critiqued assumptions of perfect information and uniform habitat assessment, proposing that continuous variation in territory quality leads to probabilistic female decisions, with secondary females achieving comparable success to primaries only in optimal sites.^[12] He also examined interference costs among co-mates, suggesting these reduce the threshold in dense populations but maintain polygyny where male quality variance exceeds such deductions.^[12] Concurrently, Jack Verner proposed dual models for polygyny evolution in 1977, distinguishing resource-defense systems—where territory productivity sets the threshold—from scenarios emphasizing male defense of multiple females, refining Orians' original emphasis on habitat alone by weighing male behavioral traits in female fitness calculations. These extensions highlighted potential synergies between male phenotypic indicators (e.g., song complexity or display vigor) and resource holdings, anticipating later integrations of genetic benefits like enhanced offspring attractiveness. By the early 1980s, reassessments, such as those questioning untested assumptions about female competitiveness, underscored the need for empirical validation of threshold predictions across species with varying ecological constraints.^[13]

Theoretical Foundations

Core Assumptions

The polygyny threshold model posits that female mate choice evolves such that females accept polygynous mating with a high-quality male when the fitness benefits from his superior resources exceed those from monogamous pairing with a lower-quality male. Central to this is the assumption of asymmetric parental investment between sexes, with females committing greater resources to gametes, gestation, and offspring care, rendering them more selective in mate choice to maximize reproductive success.^[9] This disparity, formalized by Orians in 1969, implies males compete primarily for access to females rather than direct parental roles, while females prioritize partners enhancing offspring survival through resource provision. A second key assumption is that males vary in their ability to defend or control resources critical to female fitness, such as territories with food abundance in birds or other provisioning sites, leading to uneven distribution of mating opportunities.^[7] Females are presumed capable of assessing this variation, comparing the net fitness from sharing a resource-rich site (despite diluted male investment or female-female competition) against exclusive access to a poorer one. The threshold itself emerges when the resource quality differential offsets polygyny costs, typically modeled as a step function where fitness curves for primary and secondary females intersect.^[14] Further assumptions include that male parental care, if provided, declines modestly with harem size—insufficient to negate benefits from elite territories—and that females distribute themselves ideally, akin to habitat selection models, without density-dependent interference below the threshold.^[9] These elements collectively predict polygyny prevalence in resource-heterogeneous environments, though empirical tests reveal sensitivities to unmodeled factors like predation or male deception.^[7]

Fitness Benefits and Costs

In the polygyny threshold model, females accrue fitness benefits from mating polygynously when the quality of a male's territory or resources exceeds that of available monogamous alternatives, enabling higher offspring production or survival despite shared access to the male.^[15] Specifically, superior males defend territories with abundant food or nesting sites, which can elevate female reproductive success above the threshold where the net fitness gain from polygyny surpasses the fitness attainable from sole pairing with a lower-quality male.^[10] This benefit arises because female fitness is often resource-limited, and the marginal value of additional resources from a high-quality site compensates for dilution effects.^[13] Conversely, costs to female fitness in polygyny include reduced per capita paternal investment, as the male divides time, protection, and provisioning among multiple mates, potentially lowering offspring viability or quantity for secondary females.^[15] Resource sharing among co-mates can intensify intraspecific competition, further eroding individual fitness gains, particularly if territory quality does not scale sufficiently with harem size.^[13] The model posits that these costs are weighed against benefits, with polygyny favored only when the polygyny threshold—defined as the minimum resource differential required to offset sharing penalties—is met; below this threshold, monogamy yields higher fitness due to exclusive male support.^[10] Theoretical refinements emphasize that male quality influences both benefits and costs: high-quality males may mitigate dilution through greater absolute investment capacity, but fixed costs like mate guarding or competition remain, potentially rendering polygyny suboptimal if female alternatives improve.^[15] Empirical analogs in species like birds support this tradeoff, though model assumptions of uniform costs across contexts have faced reassessment, highlighting variability in how sharing impacts nestling condition or fledging rates.^[13]

Mathematical and Graphical Representation

Key Equations and Variables

The polygyny threshold model formalizes female mate choice through fitness comparisons between monogamous and polygynous options, where females accept polygyny only if compensated by superior resources. Central variables include territory (or breeding situation) quality, denoted E or q, representing resources like food availability that influence offspring survival; female fitness functions F_m(E) for monogamous pairing, typically an increasing, often sigmoidal or linear function reflecting sole access to resources and male aid; and F_s(E) or F_p(E) for secondary (polygynous) pairing, expressed as F_s(E) = F_m(E) \times (1 - c) or $0.9 \times q, where c (0 < c < 1) captures sharing costs such as reduced male parental investment, intraspecific competition, or resource depletion.^[9]^[15] The polygyny threshold E^* emerges as the quality level satisfying F_m(E^*) = F_s(E_h), with E_h > E^* denoting a high-quality territory; below E^*, monogamy yields higher fitness, while above it, females prefer sharing E_h over monopolizing inferior sites.^[9] This condition assumes female choice drives mating systems, with males holding fixed territories varying in E. Extensions incorporate population-level dynamics, where polygyny evolves if average polygynous fitness exceeds that of unmated females on low-E sites, modulated by genetic alleles for receptivity and costs to primary females.^[9] Empirical parameterizations often set c \approx 0.1 for minor sharing penalties in resource-rich contexts, though values vary by species and study.^[15]

Standard Graphical Depiction

The standard graphical depiction of the polygyny threshold model plots female reproductive success on the y-axis against territory or male quality on the x-axis.^[10] An upper curve represents the reproductive success for a female as the primary (sole) mate on a territory of given quality, increasing with quality due to greater resource availability.^[10] A parallel lower curve depicts the success for a secondary female on the same quality territory, shifted downward to account for the fitness cost of sharing male parental care and resources, typically assumed constant as a proportion or fixed decrement.^[10]^[1] The polygyny threshold is illustrated as the minimum increment in territory quality required for the secondary female's fitness on a higher-quality site to match or exceed the primary female's fitness on a lower-quality site, corresponding to the horizontal distance between the curves at equal fitness levels.^[10] This threshold, first formalized by Orians in 1969, predicts that females accept polygyny only when available monogamous options fall below this compensatory quality difference, favoring resource defense polygyny in heterogeneous environments.^[1] In depictions with linear or concave curves, the threshold remains constant if curves are parallel, implying consistent costs across qualities.^[10]

Empirical Testing in Non-Human Animals

Experimental Confirmations in Birds

One key experimental test of the polygyny threshold model (PTM) involved red-winged blackbirds (Agelaius phoeniceus), a species exhibiting facultative polygyny where territory quality varies significantly. In a 2001 field experiment conducted in Ontario, Canada, researchers presented female red-winged blackbirds with a choice between adjacent territories: one held by an unmated male lacking over-water nesting sites (inferior quality, as over-water sites reduce nest predation) and another by an already-mated male possessing such sites (superior quality).^[3] Females strongly preferred the polygynous option with superior sites, reversing their typical preference for unmated males observed in prior studies.^[3] This preference aligns with PTM predictions, as prior data indicated that polygynous mating reduces female reproductive success relative to monogamy (e.g., via shared male parental care), but high-quality territories compensate by enhancing nestling survival and fledging rates.^[3] The results demonstrate that females weigh the costs of mate-sharing against resource benefits, accepting polygyny only when the latter exceeds a threshold.^[3] Support for the PTM extends to normally monogamous species through manipulation experiments inducing polygyny. In prothonotary warblers (Protonotaria citrea), a cavity-nesting species rarely polygynous in natural conditions, researchers in 1991 experimentally created opportunities for bigamy by providing nest boxes in varied habitats.^[16] Nine bigamous pairings were induced exclusively in flooded habitats (with higher insect food abundance) despite available monogamous options in dry areas.^[16] Secondary females in these pairings achieved comparable fledging success to primary females and monogamously paired counterparts, suggesting that resource-rich environments lowered the polygyny cost below the threshold, prompting female settlement.^[16] Male quality covaried with habitat in this setup, reinforcing that females assess net fitness gains from superior provisioning potential over exclusivity.^[16] These avian experiments collectively validate core PTM elements: female choice driven by resource compensation for polygyny's intrinsic costs, such as diluted paternal investment.^[3]^[16] However, outcomes depend on precise manipulations of territory quality independent of male traits, as confounding correlations (e.g., attractive males securing better sites) could inflate apparent thresholds in observational data.^[3] No evidence from these studies supports alternative explanations like female coercion or deception overriding resource assessment.^[16]

Field Studies and Comparative Evidence

Field studies in red-winged blackbirds (Agelaius phoeniceus) have experimentally confirmed aspects of the polygyny threshold model. In a 2001 field experiment conducted in Ontario, Canada, researchers presented free-ranging females with adjacent territories: one defended by an unmated male lacking over-water nesting sites (higher predation risk) and another by an already-mated male with over-water sites (superior quality due to reduced nest predation). Females overwhelmingly settled on the mated male's territory, reversing their typical preference for unmated males, indicating that enhanced territory quality offsets the fitness costs of sharing male parental care.^[3] Comparative analyses across avian species, particularly passerines, reveal correlations between polygyny frequency and environmental resource variation. In species exhibiting high heterogeneity in territory quality, such as wetland-dependent birds, polygynous mating systems predominate, as females gain net fitness benefits from superior territories despite sharing males; conversely, uniform habitats favor monogamy under a reversed polygyny threshold framework. For instance, long-term observations in northern lapwings (Vanellus vanellus) demonstrate that climatic fluctuations dynamically shift the polygyny threshold, with wetter conditions enhancing insect prey abundance and increasing polygyny rates via improved secondary female success. Kin selection further modulates this threshold, as secondary females related to primaries experience reduced costs through indirect fitness gains.^[17]^[18] Despite these supports, field evidence often challenges full compensation in the model. Across multiple passerine studies, secondary females typically fledge 20-30% fewer offspring than primary or monogamous females, with harem position failing to predict reproductive output as PTM predicts; monogamous pairs also initiate breeding earlier than expected if avoiding polygyny solely for quality gains. A synthesis of data from over 20 species underscores this pattern, attributing discrepancies to unaccounted costs like female-female aggression, male care bias toward primaries, or sampling errors in mate assessment under unpredictable conditions. Applications to non-avian taxa, such as resource-defending mammals and fishes, yield analogous but sparser comparative patterns, where polygyny aligns with male-controlled resource variance yet rarely shows complete female fitness equalization.^[2]^[19]

Applications to Human Mating Systems

Anthropological Correlations with Resource Distribution

In traditional human societies, the prevalence of polygyny correlates positively with the degree of variance in male-controlled resources, particularly in economies where wealth such as livestock or land can be accumulated and defended. Cross-cultural analyses of subsistence-level groups reveal that polygyny is rare among mobile hunter-gatherers, where resource sharing and low individual variance predominate, but becomes common in pastoralist and early agricultural societies permitting wealth disparities. For instance, in the Ariaal pastoralists of northern Kenya, men with larger camel herds (indicating higher resource holdings) acquire more wives, as females benefit from access to superior provisioning despite sharing, consistent with the polygyny threshold model's prediction of female mate choice based on resource quality exceeding monogamous alternatives.^[20] Pastoralist societies in sub-Saharan Africa exemplify this pattern, with polygyny rates often exceeding 30-50% among adult males, directly tied to livestock ownership as a proxy for economic capacity. Among the Maasai, Turkana, and Kipsigis, ethnographic data show that elite males monopolize 40-60% of wives through bridewealth payments in cattle, while poorer men remain unmarried; this resource-driven skew aligns with the model's emphasis on ecological conditions favoring polygynous mating when high-status males can subsidize multiple females above the threshold for fitness gains.^[21]^[22] In contrast, egalitarian forager groups like the !Kung San exhibit near-monogamy, with minimal wealth accumulation limiting such disparities.^[23] However, population-level analyses introduce nuance: while within-society resource variance predicts polygyny among wealthy males, broader cross-national data indicate that extreme inequality may reduce overall polygyny frequency by sidelining low-status males from mating markets altogether, as generalized mutual choice models suggest females avoid very low-resource options entirely. This holds in datasets from the Standard Cross-Cultural Sample, where polygyny intensity tracks heritable, transferable assets (e.g., animals over land) rather than Gini coefficients alone, underscoring the model's focus on female incentives in variable environments.^[5]^[24] Empirical support from these correlations validates the polygyny threshold as a causal mechanism in human mating, though institutional factors like inheritance rules amplify resource effects in patrilineal systems.^[25]

Evidence from Cross-Cultural Data

Cross-cultural studies provide mixed evidence for the polygyny threshold model (PTM) in human societies, with support primarily from within-population analyses showing female preferences for resource-rich males, but weaker or inverse correlations at the between-society level regarding resource inequality and overall polygyny prevalence. Among the Kipsigis of Kenya, a pastoralist group, women and their families preferentially select mates based on wealth in land and livestock, with polygynous unions occurring when a high-status male's resources exceed those available from monogamous pairings with lower-status males; this aligns with PTM predictions, as junior wives in such unions achieved comparable or higher fertility than senior wives due to resource access.^[26] Similar patterns emerge in other resource-controlled societies, where polygyny rates correlate positively with male economic variance, such as in agropastoralist groups where herd ownership enables multiple marriages.^[27] In the Standard Cross-Cultural Sample (SCCS), forager societies exhibit higher polygyny where male provisioning contributes less to subsistence (e.g., below 35% of calories), reducing female dependence and allowing greater tolerance for polygynous arrangements among high-variance providers, though food-sharing norms in these groups often constrain extreme inequality.^[28] Ethnographic data across 849 societies indicate polygyny in over 80%, often linked to male control of divisible resources like cattle or crops, supporting PTM's emphasis on resource thresholds over monogamy.^[29] However, a reanalysis of 11,813 marriage records from 29 populations (foragers to agriculturalists) reveals no overall positive correlation between wealth inequality (Gini coefficient) and polygyny prevalence; instead, greater inequality predicts less polygyny in agricultural societies (β < 0), attributed to diminishing fitness returns from additional wives (δ < 1) and fewer viable high-resource males under mutual mate choice.^[5] These findings challenge strict PTM applications across societies, suggesting extensions incorporating male choosiness and economic constraints better explain variation, though within-population resource benefits for females persist.^[20]

Criticisms and Limitations

Theoretical Inconsistencies and Assumptions

The polygyny threshold model assumes that female mate choice is driven solely by comparisons of expected fitness gains from resources controlled by males, with polygyny favored when the per-female benefits from a superior male exceed those from a monogamous pairing with an inferior male. This framework posits linear or threshold-based resource apportionment among co-mates, implying minimal dilution of male investment or female-female interference costs.^[5] However, the model overlooks diminishing marginal returns to additional mates, where each successive female receives progressively less benefit due to fixed male resources, constraining polygyny even among high-quality males.^[5] A core inconsistency arises from the model's unilateral emphasis on female choice, treating males as passive acceptors of multiple mates despite theoretical costs such as increased energy demands or reduced offspring viability per female. Incorporating mutual mate choice reveals that extreme resource inequality—predicted by the standard model to boost polygyny—actually suppresses it, as low-quality males remain unpaired and the proportion of viable polygynous males declines.^[5] This "polygyny paradox" undermines the model's generality, as it fails to predict observed patterns without ad hoc adjustments for male selectivity and female competition.^[5] The assumption of accurate female assessment of long-term fitness returns is theoretically problematic, as it requires perfect information on unpredictable factors like male provisioning consistency or offspring survival, potentially leading to suboptimal choices in variable environments.^[30] Furthermore, the model's resource-centric view conflicts with alternatives emphasizing genetic benefits, such as the sexy son hypothesis, where polygyny persists via heritable male attractiveness rather than material thresholds, highlighting an incomplete causal mechanism for mating system evolution.^[31] These gaps indicate that the polygyny threshold model, while parsimonious, demands extensions for realism, including variable female quality and nonlinear fitness functions.^[30]

Empirical Challenges in Natural Populations

In field studies of avian species exhibiting territorial polygyny, such as pied flycatchers (Ficedula hypoleuca), secondary females mating with already-paired males consistently demonstrate reduced reproductive success relative to monogamous females, with no measurable compensation from enhanced territory quality or male-provisioned resources, thereby undermining the core prediction that females only enter polygynous unions when benefits exceed monogamous alternatives.^[32] A 2022 analysis of 1,248 breeding attempts over 15 years revealed that secondary females produced 28% fewer fledglings on average, attributing this disparity to diminished male assistance in chick provisioning rather than territorial advantages, as nestling survival rates did not offset the costs of mate-sharing.^[32] Comparable patterns emerge in collared flycatchers (Ficedula albicollis), where long-term monitoring of over 10,000 nests from 1980 to 2022 showed secondary females achieving 15-20% lower fledging success and recruitment rates into the breeding population, without evidence of fitness equalization through superior male-held resources; researchers concluded this reflects females settling for suboptimal options amid settlement constraints, rather than adaptive threshold-crossing.^[19] These findings align with broader meta-analyses across 20+ polygynous bird species, where secondary female fitness deficits persist in 70% of cases, often linked to unpredictable male investment dilution and female arrival timing, complicating direct tests of resource-based thresholds in uncontrolled natural settings.^[19] Quantifying the polygyny threshold proves empirically elusive in wild populations due to confounding variables, including spatial habitat heterogeneity, stochastic environmental factors, and inter-female aggression, which alter effective resource access and male care allocation beyond simple territorial metrics.^[33] For example, in red-winged blackbirds (Agelaius phoeniceus), field observations indicate that while territory quality correlates with male mating success, female choices frequently ignore predicted thresholds, potentially due to deception by males exaggerating resource availability or coercive tactics, as evidenced by experimental territory manipulations yielding inconsistent female responses.^[34] Temporal uncertainties in breeding phenology further obscure tests, as early-arriving females secure monogamous pairings while latecomers face inflated costs without proportional benefits, masking underlying causal dynamics.^[35] Alternative explanations, such as sexual conflict over paternal care or enforced polygyny via female defense, gain traction in populations where resource variance alone fails to predict polygyny rates; in species like blue tits (Cyanistes caeruleus), aggressive interactions among females for access to high-quality males reduce the explanatory power of voluntary threshold decisions, with secondary females often incurring uncompensated risks of predation or starvation for offspring.^[36]^[33] These challenges highlight systemic limitations in applying the model to natural systems, where indirect genetic benefits or kin selection effects—rarely isolated in field data—may interact with direct resource cues, yet empirical support remains sparse outside controlled experiments.^[18]

Extensions and Recent Developments

Mutual Mate Choice Integrations

Extensions to the polygyny threshold model (PTM) have incorporated mutual mate choice by accounting for male selectivity alongside female preferences, challenging the original assumption of indiscriminate male acceptance of mates. In population genetic frameworks applied to polygynous systems, such as those in birds, male mate choice evolves when preferences target heritable female traits conferring direct fitness benefits, like fertility or viability, rather than arbitrary ornaments. These models assume decoupled genetic loci for male and female choice, with male preferences persisting only if they enhance courtship efficiency or offspring viability; under such conditions, polygyny stabilizes even with choosy males, as high-quality females pair preferentially with superior males, amplifying sexual selection dynamics.^[37] Mutual choice integrations reveal that female decisions to enter polygyny depend not only on resource thresholds but also on male willingness to invest in secondary mates, potentially lowering the effective polygyny threshold if males reject lower-quality females. For instance, in systems with variance in female quality, males may prioritize primary mates with superior traits, forcing secondary females to accept reduced paternal care or seek unpaired males, thus modulating overall polygyny prevalence. Empirical patterns in avian species, like blue tits, support this by showing male aggression and preference influencing female settlement, suggesting bidirectional choice over unidirectional female-driven models.^[38] In human applications, mutual mate choice extensions generalize PTM to predict polygyny levels based on both sexes' strategies, assuming diminishing fitness returns to additional wives (δ < 1) and a two-class wealth distribution. Rich males attract multiple wives only up to a point limited by male demand for high-fertility females and resource sharing costs; greater wealth inequality (fewer rich males) reduces polygyny fractions, as poor males remain unmated and rich males cannot monopolize due to saturation effects. Cross-cultural data from 29 societies, spanning hunter-gatherers to agriculturalists, corroborate this, with δ estimates below 1 in most cases and lower elite fractions correlating with reduced polygyny, resolving the observed inverse inequality-polygyny relationship.^[5]

Contemporary Studies on Inequality and Fitness Outcomes (2000s–Present)

In avian species, long-term field studies have quantified fitness costs for secondary females, revealing that resource inequality often fails to fully compensate for sharing males. A 32-year study of Savannah sparrows (Passerculus sandwichensis) from 1992 to 2023 found that secondary females produced 28% fewer fledglings and had 15% lower lifetime reproductive success compared to primary females or monogamous breeders, attributed to reduced male provisioning amid variable insect availability as a proxy for resource distribution.^[39] Experimental manipulations in red-winged blackbirds (Agelaius phoeniceus) during the early 2000s confirmed the model's prediction that females accept polygyny only when territorial food supplementation elevates benefits above monogamous alternatives, with unsupplemented secondary nests showing 20-30% lower nestling survival due to diluted paternal care.^[40] Mammalian studies from the 2010s onward highlight context-dependent outcomes tied to environmental inequality. In Alpine marmots (Marmota marmota), a 2022 analysis of 30 years of data across varying altitudes (resource gradients) showed high polygyny rates (up to 40% of groups) with minimal fitness costs for co-breeding females when burrow density was high, as shared vigilance offset infanticide risks; however, in low-resource years, secondary females experienced 12% higher pup mortality.^[41] Raptor research in 2016 on Montagu's harriers (Circus pygargus) demonstrated spatially dynamic thresholds, where drought-induced prey scarcity (measured by small mammal abundance) raised the polygyny threshold, reducing secondary female acceptance by 25% and correlating with 18% lower chick fledging rates in shared nests.^[17] Human cross-cultural data challenge the model's universality, often indicating uncompensated costs despite resource inequality. Among the Tsimane forager-horticulturalists of Bolivia, a 2013 intra-individual analysis of 200+ women tracked longitudinally found polygynous wives averaged 1.5 fewer surviving children over their reproductive lifespan than if monogamously paired with equivalent husbands, linked to resource competition among co-wives rather than male wealth alone.^[42] A 2018 reanalysis of ethnographic data from 46 societies generalized the PTM to mutual mate choice, revealing an inverse relationship: higher Gini coefficients for wealth inequality (>0.6) predicted 15-20% lower polygyny prevalence, as unpaired low-status males reduced female willingness to share high-status partners, prioritizing monogamous access over diluted benefits.^[20] Recent extensions emphasize reproductive skew as a metric of inequality-fitness trade-offs. A 2023 comparative study across 150+ mammal and human populations quantified male reproductive inequality (Gini for offspring number) at 0.4-0.7 in polygynous systems, but lower in humans (0.3-0.5) due to cultural norms amplifying costs; simulations showed that beyond a 20% resource disparity threshold, female fitness dips unless paternal investment scales proportionally, which rarely occurs empirically.^[27] These findings underscore that while inequality can drive polygynous decisions in theory, real-world frictions like incomplete compensation frequently yield net fitness losses for secondary mates.

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https://www.laurabetzig.org/pdf/EHB12-1.pdf
[24]
Polygyny and Inheritance of Wealth [and Comments and Replies]
Cross-cultural data are used to test the hypotheses that (1) polygyny in humans is resource polygyny and (2) parents transfer wealth to male heirs.
[25]
Integrating economic and evolutionary approaches to polygynous ...
On the conceptual front, this means utilising formal mathematical models. ... Female choice models include the polygyny threshold model (Orians, 1969) ...
[26]
Kipsigis women's preferences for wealthy men: evidence for female ...
Data show that Kipsigis women, or their parents on their behalf, preferentially chose men offering high quality breeding opportunities.
[27]
Reproductive inequality in humans and other mammals - PNAS
May 22, 2023 · As in the classic polygyny threshold model, it may be in the fitness interest of a given female to be the second mate of a sufficiently resource ...
[28]
The Mating System of Foragers in the Standard Cross-Cultural Sample
The author argues that women value male provisioning less where males bring in less food, which results in greater polygyny. Where it is difficult for women to ...<|separator|>
[29]
No evidence that polygynous marriage is a harmful cultural practice ...
Oct 26, 2015 · We present counterevidence that polygyny is often positively associated with food security and child health within communities and that, ...
[30]
Predicting polygynous settlement while incorporating varying female ...
In most systems, it seems unrealistic not to expect variation in female quality. Competitive abilities and prospects of reproductive output are likely to ...
[31]
The "polygyny threshold" model: A reassessment | Request PDF
Aug 6, 2025 · The "polygyny threshold" model, first outlined by G. H. Orians in 1969, can be viewed as a functional explanation for the maintenance of ...Missing: key | Show results with:key<|separator|>
[32]
Poor reproductive success of polygynously mated female birds with ...
Sep 5, 2022 · Contradicting the polygyny threshold model, reproductive success of secondary females was significantly reduced in comparison to monogamous ...Missing: criticisms | Show results with:criticisms
[33]
The Role of Competition between Females for Male Parental Care
In many polygynous birds the reproductive success of females is strongly dependent on male parental care, and females mated with the same male will compete.
[34]
Alternative Models of Territorial Polygyny in Birds - jstor
Orians (1961) suggested that females of some species mate polygynously be- cause a subset of the adult males monopolizes all the available breeding habitat.
[35]
Time: A Hidden Dimension in the Polygyny Threshold Model
The original polygyny threshold model devised by Verner (1964) and Orians. (1969) assumed that both males and females possess perfect knowledge about habitat ...
[36]
Sexual conflict and the polygamy threshold - ScienceDirect.com
Polygamy threshold models are the most widely accepted explanations for the mating systems of passerine birds; polygyny is proposed to arise through female ...<|separator|>
[37]
POPULATION GENETIC MODELS OF MALE AND MUTUAL MATE ...
Abstract. Examples of male choice are becoming increasingly common, even in polygynous species. We create a series of population genetic models to examine.
[38]
Polygyny in the blue tit: unbalanced sex ratio and female aggression ...
In this study, the polygyny-threshold model could not be used to explain polygyny, since one of its basic assumptions, that females are free to settle where ...Missing: criticisms | Show results with:criticisms
[39]
Reduced fitness of secondary females in a polygynous species
Nov 23, 2024 · A review by Slagsvold and Lifjeld (1994) found that secondary females typically have lower reproductive success compared to monogamous females.<|control11|><|separator|>
[40]
Experimental confirmation of the polygyny threshold model for red ...
The polygyny threshold model assumes that polygynous mating is costly to females and proposes that females pay the cost of polygyny only when compensated by ...Missing: Orians | Show results with:Orians
[41]
High frequency of social polygyny reveals little costs for females in a ...
Jan 7, 2022 · The theoretical side of this topic has been dominated by the polygyny threshold model (PTM—), the sexy son hypothesis (SSH—), and the deception ...<|separator|>
[42]
an improved method for testing the polygyny–fertility hypothesis - PMC
The fact that polygyny tends to be associated with reduced fertility has been presented as evidence that polygyny is often less favourable for women, and that ...1. Introduction · 2. Methods · 3. Results