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Evolution; International Journal Of Organic Evolution[JOURNAL]

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Natural and sexual selection drive the evolution of carotenoid-based bare-part ornamentation in shorebirds.

Minias P, Kamiński M, Janiszewski T

Evolution · 2026 May · PMID 41805985 · Publisher ↗

Carotenoid-based plumage ornaments in birds have long been acknowledged to yield honest signal of individual quality and play a role in intra- and inter-sexual interactions. However, much less scientific attention has be... Carotenoid-based plumage ornaments in birds have long been acknowledged to yield honest signal of individual quality and play a role in intra- and inter-sexual interactions. However, much less scientific attention has been devoted to the role and evolution of non-plumage (bare-part) avian ornamentation. Here, we aimed to investigate processes that shaped the evolution of carotenoid-based ornamentation of bare parts (bill and legs) in Charadrii and Scolopaci shorebirds. Our phylogenetically informed comparative analysis across all extant shorebirds revealed that both natural and sexual selection contributed to the evolution of carotenoid-based bare-part ornamentation. We found evolutionary associations between carotenoid pigmentation and habitat variation, climate, and body size, advocating for the importance of ecological, environmental, and life history traits. Sexual selection (mating system bias and sexual size dimorphism) was also identified as an important driver in the evolution of carotenoid-based ornamentation. Underrepresentation of combinations of carotenoid- and melanin-based ornaments suggested overlapping functions of ornamental traits with different developmental origins or excessive costs associated with their simultaneous expression. Our study adds to the understanding of the complex and multi-faceted processes responsible for the evolution of remarkably diverse avian ornamentation.

Extreme sexual size dimorphism correlates with proxies of postcopulatory sexual selection across selected Dolomedes fishing spiders.

Linde M, Donke E, Schwartz SK … +2 more , Hebets EA, Michalik P

Evolution · 2026 May · PMID 41805983 · Publisher ↗

Sexual size dimorphism (SSD) and mating system diversity are key factors shaping reproductive evolution, yet their relationships with post-mating sexual selection remain incompletely understood. Here, we use a comparativ... Sexual size dimorphism (SSD) and mating system diversity are key factors shaping reproductive evolution, yet their relationships with post-mating sexual selection remain incompletely understood. Here, we use a comparative framework across multiple species of Dolomedes fishing spiders to examine how mating systems, sperm production strategies, and proxies of postcopulatory selection relate to SSD. We quantified testes size and spermatogenesis in three closely related species-D. tenebrosus, D. okefinokensis, and D. scriptus-that differ in mating system and SSD. Dolomedes tenebrosus, a monogynous species with extreme female-biased SSD (eSSD) and male self-sacrifice, showed evidence of termination of spermatogenesis at maturity, indicating a terminal investment strategy. Dolomedes okefinokensis exhibited a similar pattern, suggesting convergent evolution toward monogyny. In contrast, D. scriptus, with moderate SSD and a polygynous system, maintained continuous spermatogenesis, consistent with repeated mating opportunities. Expanding to 13 Dolomedes species, we found a strong positive association between eSSD, monogyny, and spermatogenesis termination, alongside increased spermophor volume in eSSD species. These findings suggest that eSSD is associated with both a terminal male reproductive strategy and intensified investment in sperm transfer structures. Together, our results highlight an integrated evolutionary link between SSD, mating system, and male reproductive investment across Dolomedes spiders.

Gene frequency changes and a per-generation time-scale for continuous-time populations with arbitrary size variations.

Hössjer O, Ewens W

Evolution · 2026 May · PMID 41802202 · Publisher ↗

One of the most fundamental calculations needed in an examination of the effect of natural selection in a Mendelian population is to find the rate of change in continuous-time of the frequency of any allele under natural... One of the most fundamental calculations needed in an examination of the effect of natural selection in a Mendelian population is to find the rate of change in continuous-time of the frequency of any allele under natural selection, and from this to find the number of generations needed for the frequency of an allele to change from one given value to another. In this paper, we analyze the presently accepted self-contained continuous-time theory of allele frequency change for large populations caused only by natural selection, which assumes no mutation and no gene frequency changes due to random genetic drift in a monoecious diploid population. We claim that the present theory is incomplete, with severe restrictions on how the population size varies over time, and with no natural generations concept built in. In order to remedy this, we propose a new self-contained theory, making throughout the same assumptions described above concerning mutation and random genetic drift, and also assuming a monoecious diploid population.

The double-edged effect of environmental fluctuations on evolutionary rescue.

Shibasaki S, Yamamichi M

Evolution · 2026 Jun · PMID 41790495 · Publisher ↗

Recent studies revealed that contemporary evolution can prevent population extinction in deteriorating environments. Such evolutionary rescue has been intensively studied, but few have focused on environmental fluctuatio... Recent studies revealed that contemporary evolution can prevent population extinction in deteriorating environments. Such evolutionary rescue has been intensively studied, but few have focused on environmental fluctuations. As global changes alter both the mean and variance of environmental variables, it is crucial to understand how environmental fluctuations affect evolutionary rescue. Here, through the evolution experiments on green algae Chlorella vulgaris, we show that increasing the amplitude of environmental fluctuations around long-term deteriorating trends has negative and positive effects on evolutionary rescue. We first increased the salinity level gradually to 0.6M NaCl and found that the algae exposed to large fluctuations tended to grow more slowly. This seems to be because large fluctuations produce an episode of a huge environmental change, which can increase adaptation lag. Then, we increased the salinity level to 1M NaCl and found that the algae exposed to large fluctuations grew, while those exposed to smaller or no fluctuations did not. This seems enigmatic, but our mathematical model suggests that trait variance within a population might increase under large fluctuations, which can promote adaptive evolution. Our results highlight the complex role of environmental fluctuations in evolutionary rescue, calling for more investigations to understand evolutionary rescue in nature.

How is variation in fitness maintained?

Charlesworth B

Evolution · 2026 May · PMID 41790487 · Publisher ↗

The existence of substantial additive genetic variance in fitness poses a long-standing question for evolutionary geneticists. A new theoretical paper by Connallon and Czuppon examines the effect of random genetic drift... The existence of substantial additive genetic variance in fitness poses a long-standing question for evolutionary geneticists. A new theoretical paper by Connallon and Czuppon examines the effect of random genetic drift and temporal fluctuations in fitness on loci maintained by balancing selection. It suggests ways in which substantial additive variance can be maintained, by causing loci to depart from their equilibria under selection alone.

Wing length canalisation and behaviour across birds: a phylogenetic meta-analysis of variance.

Reinhold K, Sánchez-Tójar A

Evolution · 2026 Jun · PMID 41773955 · Publisher ↗

We examined the expectation that stronger stabilising selection leads to a decrease in trait variation across species by investigating individual variation in wing length. We hypothesised that species that heavily rely o... We examined the expectation that stronger stabilising selection leads to a decrease in trait variation across species by investigating individual variation in wing length. We hypothesised that species that heavily rely on aerial feeding, as well as long-distance migratory species, show higher canalisation (lower coefficients of variation, i.e., CV values) in wing length than non-aerial feeders and non-migratory species. We collected summary statistics on wing length for males and females from the literature and analysed them using recently developed meta-analytic metrics for integrating phenotypic variance estimates. Our phylogenetic multilevel meta-analysis showed relatively low heterogeneity among CV values, indicating generalisability of the overall CV value (2.6%). Although not all pairwise comparisons were statistically significant, all our analyses consistently showed higher canalisation in aerial compared to non-aerial feeders, and in migratory compared to non-migratory species. We conclude that wing length in bird species relying on their wings more heavily is likely under stronger (stabilising) selection, which in turn would have led to the observed higher canalisation on this trait for those species. Our study showcases how to combine already available descriptive statistics for phenotypic traits with underused meta-analysis of variance approaches to test often-neglected evolutionary predictions.

Pollinator assemblage composition predicts trait divergence in a pollination-generalized plant.

Torres-Vanegas F, Temesvári V, García Y … +2 more , Friberg M, Opedal ØH

Evolution · 2026 May · PMID 41757704 · Publisher ↗

The causal role of pollinators in driving the divergence of plant traits is a fundamental tenet of angiosperm evolution, providing hallmark examples of evolution in response to natural selection. However, it remains uncl... The causal role of pollinators in driving the divergence of plant traits is a fundamental tenet of angiosperm evolution, providing hallmark examples of evolution in response to natural selection. However, it remains unclear how geographic variation in pollinator assemblages relates to the divergence of pollination traits in pollination-generalized plants. We characterized pollinator assemblages that interacted with Viscaria vulgaris in southern Sweden, and evaluated, through statistical dimension reduction, whether pollination traits were associated with an inferred main axis of geographic variation in pollinator assemblages. We documented a functionally broad range of pollinators that visited V. vulgaris. Although the most frequent pollinator functional groups were present in most populations, their relative contribution to flower visitation varied across the study area, establishing a geographic mosaic of pollinator assemblages. We demonstrate that the geographic variation in pollinator assemblages can predict the divergence of pollination traits in V. vulgaris. The findings of this geographic comparative study are consistent with the hypothesis that geographic variation in pollinator assemblages drives the divergence of pollination traits in pollination-generalized plants. Thus, generalized plant-pollinator interactions do not preclude the divergence of pollination traits, which may maximize the collective contribution of local pollinator assemblages rather than that of a principal pollinator.

Caenorhabditis diversity on Pohnpei, Micronesia, provides evidence that the Elegans Supergroup has its roots in the Americas and diversified in the Pacific en route to Asia.

Rockman MV, Tintori SC, Nguyen THM … +1 more , Yomai VMH

Evolution · 2026 May · PMID 41757685 · Publisher ↗

Model organisms are powerful tools for discovery in cell and molecular biology, and studies of their natural history have the potential to provide bridges between these fields and ecology and evolutionary biology. The ne... Model organisms are powerful tools for discovery in cell and molecular biology, and studies of their natural history have the potential to provide bridges between these fields and ecology and evolutionary biology. The nematode Caenorhabditis elegans is a preeminent model, and recent findings place its center of diversity in the cool, high-elevation forests of Hawaii. To test models of biogeography and species coexistence, we investigated Caenorhabditis on Pohnpei, an island in Micronesia, home to the largest patch of high-elevation forest between Hawaii and Asia. We found nine species of Caenorhabditis, five of them new. Using the distribution of nematodes among habitat patches, we parameterized models of Caenorhabditis population biology that help explain species coexistence patterns. We inferred a phylogeny for 70 species of Caenorhabditis and performed the first quantitative biogeographic analysis for the group. Our analysis suggests that the deep ancestors of the Elegans Supergroup of species lived in the Americas. The Supergroup's subsequent diversification occurred in Oceania, giving rise to a diverse Oceanian fauna and ultimately to multiple lineages that moved into Asia, Africa, Australasia, and back into the Americas. We infer a slow trans-Pacific migration, with the islands of Oceania serving as sources rather than sinks for biodiversity.

Experimental evolution in communities: beyond pairwise interactions.

Montbel V, Hrcek J

Evolution · 2026 May · PMID 41746203 · Publisher ↗

Experimental evolution is a powerful method that has been instrumental for revealing core mechanisms of adaptation and coevolution. It has mostly been used in very simple settings of 1 or 2 species. Yet, it is now increa... Experimental evolution is a powerful method that has been instrumental for revealing core mechanisms of adaptation and coevolution. It has mostly been used in very simple settings of 1 or 2 species. Yet, it is now increasingly being employed in more complex community settings that include indirect effects, higher-order interactions, and multidimensional selection typical of natural communities. Here we synthesize the emerging field of experimental evolution in communities and show how community context reshapes selection and evolutionary trajectories, beyond what single-species or pairwise designs predict. We conducted a systematic literature survey targeting multi-species, multi-generation evolution, identifying 100 such studies with the number increasing recently. Despite this progress, most experiments are biased toward microbial systems and competitive interactions, leaving major gaps for predicting evolution in realistic communities. We discuss community ecology concepts in the light of experimental evolution, together with designs that address these concepts. We emphasize 3 main research areas: indirect and higher-order interactions that make selection multidimensional, eco-evolutionary feedbacks linking trait change to community dynamics, and genetic constraints that shape responses across interaction networks. We then discuss routes to increase ecological realism with field experiments and conclude by outlining key research fronts for experimental evolution in communities.

Digest: Population-level processes that facilitate divergence differ between Andean and Amazonian birds.

Gokhale P

Evolution · 2026 May · PMID 41739061 · Publisher ↗

Wacker and Winger (2026) examine population-level divergence across 2 very different but connected landscapes, the Andes and the Amazon, using genomic data from eight Tangara bird species. Andean populations show strong... Wacker and Winger (2026) examine population-level divergence across 2 very different but connected landscapes, the Andes and the Amazon, using genomic data from eight Tangara bird species. Andean populations show strong genetic structure, with lower gene flow and genetic diversity. In the Amazon, genetic structure is weak, with higher gene flow and genetic diversity. These results suggest that divergence can happen more easily in the Andes due to stronger geoclimatic barriers than in lowland Amazonia.

Addressing multi-generational non-genetic responses in experimental studies of evolution.

Zilio G, Bedhomme S, Fronhofer EA … +4 more , Jacob S, Legrand D, Philippe H, Chevin LM

Evolution · 2026 Jun · PMID 41728916 · Publisher ↗

Populations that face environmental change reducing their mean fitness can recover by adaptive genetic evolution over multiple generations, but their immediate responses may also involve non-genetic mechanisms, though th... Populations that face environmental change reducing their mean fitness can recover by adaptive genetic evolution over multiple generations, but their immediate responses may also involve non-genetic mechanisms, though the latter can be difficult to demonstrate. When the dynamics of such non-genetic changes in mean phenotype and fitness span multiple generations, their effects at the population level can be difficult to distinguish from those of natural selection on genetic variants. While the existence of non-genetic inheritance is no longer controversial, we argue that its potential contribution to observed patterns in evolutionary studies remains overlooked, especially for processes leading to phenotypic change that unfolds over multiple generations, which we call multigenerational non-genetic responses (MUNGER). We highlight three major forms of MUNGER that, if not properly accounted for, could confound inference about genetic changes: delayed impact of stress, transgenerational plasticity, and priming. We summarize how each may impact the dynamics of phenotypic change across generations in concrete experimental contexts (e.g., experimental evolution, common gardens, ecotoxicological experiments). We propose that analysing the dynamic properties of MUNGER, their relative contributions to overall phenotypic responses, and how they interact with genetic changes, should help build a more comprehensive understanding of evolutionary responses to changing environments.

The land-to-water transition impacts brain shape in caniform carnivorans.

Kirkwood TMB, Shultz S, Silcox MT … +4 more , Schwab JA, Palmer SM, Rule JP, Jones KE

Evolution · 2026 May · PMID 41700709 · Publisher ↗

Mammals exhibit remarkable diversity in brain size and morphology, resulting from numerous ecological radiations throughout the Cenozoic. Although previous studies have demonstrated the influence of phylogeny, allometry,... Mammals exhibit remarkable diversity in brain size and morphology, resulting from numerous ecological radiations throughout the Cenozoic. Although previous studies have demonstrated the influence of phylogeny, allometry, and various ecological variables on endocranial morphology in certain mammalian lineages, the extent to which locomotor habitat influences brain shape evolution remains unclear. The suborder Caniformia, or "dog-like" carnivorans, is an ecologically diverse clade, containing terrestrial, arboreal, fossorial, and semiaquatic species, each facing a unique set of sensory and motor challenges. Here, we evaluate the impact of phylogeny, allometry, and locomotor habitat on brain shape evolution in caniforms. We examine endocranial morphology in 73 species using high-density 3D geometric morphometrics, principal component analysis, and phylogenetic comparative methods. Our findings indicate that the land-to-water transition has a significant impact on endocranial shape across caniforms. This effect is more pronounced in pinnipeds than in other semiaquatic fissipeds, such as mustelids and ursids, likely due to their more derived ecology and greater commitment to aquatic life. Aquatic caniforms tend to exhibit expanded cerebra alongside reduced paleocortices and olfactory bulbs. These morphological changes likely reflect selection pressures acting on both the brain and the surrounding cranial architecture in aquatic environments.

A quantitative genetics analysis of evolution in the hominoid appendicular skeleton.

Cooper MJ, Conaway MA, von Cramon-Taubadel N

Evolution · 2026 May · PMID 41697136 · Publisher ↗

Hominoidea are a diverse superfamily, with potential selection on the postcranium from divergent locomotor patterns. The scarcity of postcranial primate fossil remains, however, means that selection on the appendicular s... Hominoidea are a diverse superfamily, with potential selection on the postcranium from divergent locomotor patterns. The scarcity of postcranial primate fossil remains, however, means that selection on the appendicular skeleton has been difficult to study directly. Here, we use complementary methods from evolutionary quantitative genetics to assess selection versus drift within hominoids and an outgroup of cercopithecoid monkeys. While most branches were characterized by drift or stabilizing selection, directional selection was indicated on the branch leading to Homo sapiens and potentially on the branch to Hylobates lar. Directional selection gradients were then assessed for each trait on these 2 lineages. Strong selection was detected on limb length and traits of the os coxa in both lineages. Our results are consistent with prior studies of evolution in the hominoid postcranium and indicate that these methods may be useful in studying multiple elements in conjunction for a holistic assessment of evolution in the primate skeleton in the absence of extensive fossil data.

Embryology reveals a morphological signature of ancestral diurnality maintained in a nocturnal lineage.

Griffing AH, Bauer AM, Wegerski AE … +2 more , Gamble T, Rasys AM

Evolution · 2026 May · PMID 41689508 · Publisher ↗

Evolutionary transitions to diurnality are often associated with specialized ocular morphology, such as pits (foveae) in the retina. Foveae are required for high visual acuity and are found primarily in diurnal vertebrat... Evolutionary transitions to diurnality are often associated with specialized ocular morphology, such as pits (foveae) in the retina. Foveae are required for high visual acuity and are found primarily in diurnal vertebrate taxa, including lizards. Geckos have undergone repeated evolutionary transitions between nocturnality and diurnality. Aristelliger is a nocturnal lineage embedded within a largely diurnal clade and represents a putative reversal to nocturnality. We investigated eye development and morphology of Aristelliger geckos. Despite sharing a most recent common ancestor with its diurnal sister lineage over 60 million years ago, Aristelliger exhibits a fovea. Fovea development is coordinated by dynamic changes in eye shape-such as ocular elongation and retraction-during embryonic development. In Aristelliger embryos, these elongation and retraction phases occur in the same region where the fovea forms, closely resembling the pattern seen in Anolis, a diurnal lizard with two foveae. Given the retention of a fovea in Aristelliger, we hypothesize that the transition to nocturnality is relatively recent. Incorporating these results into comparative phylogenetic methods results in a hypothesized diurnal ancestral temporal niche for sphaerodactylid geckos. We illustrate, similar to some nocturnal primates, that developmental data and robust morphological investigation can provide unique insights into macroevolutionary studies.

To survive in the cold: the evolution of reduced decay rate in a bacteriophage.

Chu XL, Zhang QG, Buckling A

Evolution · 2026 May · PMID 41671162 · Publisher ↗

Viruses typically have high decay rates (mortality rates outside hosts), and applications of phage viruses for combating harmful bacterial in clinical and agricultural contexts would favor slow-decaying phage materials.... Viruses typically have high decay rates (mortality rates outside hosts), and applications of phage viruses for combating harmful bacterial in clinical and agricultural contexts would favor slow-decaying phage materials. There is evidence for a trade-off between viral survival and growth rate, which may constrain the evolution of reduced decay rate. Temperature is likely to affect the optimal balance of this trade-off; e.g., faster growth may be more beneficial at warmer temperatures where phages spend less time outside of hosts in waiting for encountering a new host individual. We tested this hypothesis by experimentally evolving a lytic phage that infects the bacterium Pseudomonas fluorescens. Phages evolved at two temperatures for 20 cycles of dilution and propagation, with the ancestral bacteria being supplied every cycle. Phage populations from different temperatures showed different modes of adaptation in growth and decay traits. In particular, phages that evolved at the colder temperature showed a reduction in decay rate, regardless of assay conditions. Our results suggest phage training programs and resource collecting efforts to focus on cold environments for slow-decaying phage materials.

Discriminating models of trait evolution.

Roa Lozano J, Jangra S, DeGiorgio M … +2 more , Assis R, Adams R

Evolution · 2026 Apr · PMID 41661137 · Full text

A central challenge in comparative biology is linking present-day trait variation across species with unobserved evolutionary processes that occurred in the past. In this endeavor, phylogenetic comparative methods are in... A central challenge in comparative biology is linking present-day trait variation across species with unobserved evolutionary processes that occurred in the past. In this endeavor, phylogenetic comparative methods are invaluable for fitting, comparing, and selecting evolutionary models of varying complexity and biological meaning. Traditionally, evolutionary studies have relied on conventional statistical approaches to assess model fit and identify the one that best explains variation in a given trait. Here, we explore an alternative strategy by applying supervised learning to predict evolutionary models via discriminant analysis. We formally introduce Evolutionary Discriminant Analysis (EvoDA) as an addition to the biologist's toolkit, offering a suite of new methods for studying trait evolution. We evaluate the performance of EvoDA alongside conventional model selection through a series of fungal phylogeny case studies, each targeting increasingly challenging analytical tasks. These results showcase the strengths of EvoDA, with substantial improvements over conventional approaches when studying traits subject to measurement error, which likely reflect realistic conditions in empirical datasets. To complement our simulation-based benchmarking, we explore the application of EvoDA for tackling a notoriously difficult task: predicting the mode and tempo of gene expression evolution. This empirical analysis suggests that stabilizing selection acts on a majority of genes, with bursts of expression evolution in a handful of genes related to stress, cellular transportation, and transcription regulation. Collectively, our findings illustrate the promise of EvoDA for predicting trait models across a range of evolutionary and experimental contexts, establishing a new methodological framework for the next era of comparative research.

Latitude and communal living are associated with the evolution of joint territory defense in New World songbirds.

Arashanapalli S, Freeman BG

Evolution · 2026 Apr · PMID 41637056 · Publisher ↗

There is great variation across species in how many individuals participate in territorial defense. Here, we test the hypotheses that joint territorial defense by pairs or family groups is more common in songbird species... There is great variation across species in how many individuals participate in territorial defense. Here, we test the hypotheses that joint territorial defense by pairs or family groups is more common in songbird species that (1) nest in tropical latitudes, (2) exhibit weak sexual selection, (3) maintain long-term social bonds, (4) defend year-round territories, (5) nest cooperatively, and (6) are sedentary. We conducted the first broad-scale test of these hypotheses by performing 3,177 playback experiments across the Americas to measure territorial defense behaviors for 264 species. We found support for three of our six predictions: tropical species, cooperative nesters, and species with long-term social bonds are indeed more likely to jointly defend territories, but other variables were unrelated to joint territorial defense. Latitudinal zone was the strongest predictor, suggesting that tropical environments select for joint territory defense above and beyond the life-history traits we included in our analysis. The remaining traits that predicted territorial defense describe aspects of communal living, though the association with long-term social bonds was marginal. Overall, we document a strong latitudinal gradient wherein joint territorial defense is consistently more common in the tropics even when accounting for different life-history traits of tropical birds.

The inhibitory cascade model and evolution in segmentally organized tissues.

Auerbach BM, Roseman CC

Evolution · 2026 Jun · PMID 41637054 · Publisher ↗

The inhibitory cascade model (ICM) of morphogenesis is an effort to link development to the production of variation, which can influence evolutionary trajectories. The ICM proposes that serially developing features, such... The inhibitory cascade model (ICM) of morphogenesis is an effort to link development to the production of variation, which can influence evolutionary trajectories. The ICM proposes that serially developing features, such as molar teeth, are governed by the relative magnitudes of one activating and one inhibiting developmental process. The statistical expectations of the ICM are typically expressed and analyzed on a first-element standardized scale and seem to be a good predictor of molar proportions. However, the ICM has been applied to traits that occur in series but do not develop in sequence and still recovers as good a fit as when applied to serially developing traits. Such an undiscriminating result raises questions about whether the fit of the ICM is an artifact of standardization. The mathematical rendition of the ICM does not correspond with the verbal descriptions of the developmental argument. Applying our novel rearticulation of the ICM to biological, nonbiological, and simulated data, we demonstrate that the apparent goodness of fit of the ICM to many biological systems is an artifact of scaling correlated values with a common denominator. There is no evidence supporting the ICM at the developmental, variational, or evolutionary levels.

Is the fundamental theorem of natural selection of any use?

Charlesworth B

Evolution · 2026 Apr · PMID 41618867 · Publisher ↗

There have been many recent discussions of the Fundamental Theorem of Natural Selection, with an emphasis on its mathematical accuracy. It is argued here that, despite the mathematical problems that have been uncovered,... There have been many recent discussions of the Fundamental Theorem of Natural Selection, with an emphasis on its mathematical accuracy. It is argued here that, despite the mathematical problems that have been uncovered, it still has utility for biologists. In particular, it predicts an absence of additive genetic variance for fitness for populations at equilibrium under selection alone, a result that is valid under very general conditions. This raises the question as to why there are such high levels of additive variance in fitness and fitness components, but little evidence for nonadditive variance.
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