Maintenance Of Polymorphism Research Articles

Matings Between Individuals with Similar Major Histocompatibility Complex (MHC) Improve Offspring Survival in the Rainbow Trout (Oncorhynchus mykiss)

The major histocompatibility complex (MHC) consists of genes involved in immune response and molecular discrimination between self and non-self. MHC genes are the most polymorphic in vertebrates. The origin and maintenance of polymorphism in MHC genes in populations is still unresolved. Mechanisms such as sexual selection and heterozygote advantage have been suggested as explanations for this high variability. In this study, a farmed population of rainbow trout (Oncorhynchus mykiss) characterized by the presence of specific MHC class IIB gene haplotypes at a frequency higher (30%) than that expected from random matings was investigated. Therefore, it was hypothesized that disassortative matings occur with an adaptive advantage for females, resulting in improved reproductive performance when mated with individuals with similar MHC haplotypes. Genetic analyses of the breeders were performed to define the MHC haplotypes and to perform specific matings. The effect of mating was evaluated by analyzing the survival rate of the offspring at various stages of incubation until swim-up. The reproductive performance of the offspring derived from specimens with similar haplotypes showed a better survival trend during the first life stages and reduced malformations. The results obtained are in contrast with the heterozygous advantage theory, therefore it was hypothesized, as for other salmonid species, the presence of a positive selection towards locally adapted MHC genes that promotes reproduction between genetically similar individuals.

The Evolution of Extreme Genetic Variability in a Parasite-Resistance Complex.

Genomic regions that play a role in parasite defense are often found to be highly variable, with the MHC serving as an iconic example. Single nucleotide polymorphisms may represent only a small portion of this variability, with Indel polymorphisms and copy number variation further contributing. In extreme cases, haplotypes may no longer be recognized as orthologous. Understanding the evolution of such highly divergent regions is challenging because the most extreme variation is not visible using reference-assisted genomic approaches. Here we analyze the case of the Pasteuria Resistance Complex (PRC) in the crustacean Daphnia magna, a defense complex in the host against the common and virulent bacterium Pasteuria ramosa. Two haplotypes of this region have been previously described, with parts of it being non-homologous, and the region has been shown to be under balancing selection. Using pan-genome analysis and tree reconciliation methods to explore the evolution of the PRC and its characteristics within and between species of Daphnia and other Cladoceran species, our analysis revealed a remarkable diversity in this region even among host species, with many non-homologous hyper-divergent-haplotypes. The PRC is characterized by extensive duplication and losses of Fucosyltransferase (FuT) and Galactosyltransferase (GalT) genes that are believed to play a role in parasite defense. The PRC region can be traced back to common ancestors over 250 million years. The unique combination of an ancient resistance complex and a dynamic, hyper-divergent genomic environment presents a fascinating opportunity to investigate the role of such regions in the evolution and long-term maintenance of resistance polymorphisms. Our findings offer valuable insights into the evolutionary forces shaping disease resistance and adaptation, not only in the genus Daphnia, but potentially across the entire Cladocera class.

A field investigation into potential reproductive and resistance advantages of purple-petalled plants in polymorphic populations of horsenettle (Solanum carolinense)

Background and aims – Variation in flower color among individuals within plant populations has long fascinated horticulturalists and evolutionary biologists alike. Nevertheless, the diversity of genetic factors and ecological interactions that act to maintain flower-color polymorphisms over time are understood for very few species. Here, we suggest that Solanum carolinense (horsenettle) makes an excellent model system for studying the evolutionary ecology of a flower-color polymorphism, and we begin to shed insight into factors that might affect the stability of the polymorphism. Material and methods – We transplanted 24 horsenettle ramets of each of 40 field-collected genets (10 genets per each of four source fields) into a common-garden plot (in an oldfield with an existing horsenettle population). For each ramet, we visually characterized petal color and quantified production of flowers, fruits, seeds, and damage by 11 species of herbivores. Key results – Rather than varying continuously within ramets and among genets, petal color fell into three relatively discrete categories that are common in most horsenettle populations: purple, white, and mauve. In the common-garden experiment, purple-petalled morphs initiated significantly more flower buds, opened more flowers, and produced more fruits and seeds than did the mauve-petalled or white-petalled morphs. Petal-color morphs differed only slightly in resistance to a minority of the herbivore community, with no consistent advantage for color morphs across species of herbivores or types of tissues fed upon. Conclusion – Horsenettle has numerous characteristics that make it an excellent system for studying the genetics and ecology that drive the maintenance of flower-color polymorphisms. Although pleiotropic effects of flower-color on resistance to herbivory appear to be minimal in horsenettle, potential pleiotropies that differentially affect maternal and paternal reproduction, or sexual and vegetative reproduction, are worthy of further study.

Long‐Term Maintenance of Complex Chromosomal Inversion Polymorphism in Drosophila mediopunctata

ABSTRACTNatural selection is known to favor specific gene combinations, thereby shaping the evolution of recombination rates, often through epistatic interactions. However, the dynamics of these interacting factors within natural populations remain poorly understood. In this study, we investigate the long‐term maintenance of a complex polymorphism involving linked, nonoverlapping chromosomal inversions in a natural population of Drosophila mediopunctata. Remarkably, even after 30 years—equivalent to roughly 340 generations—two major features have remained unexpectedly stable: the linkage disequilibrium (LD) between inversions, which deviates significantly from the theoretical prediction of decay, and a consistent seasonal cycle pattern of heterozygous excess and homozygous deficiencies. We explored the roles of recombination suppression, epistatic selection, and overdominance in maintaining this stability, examining their alignment with previously described patterns. Our findings reveal that moderate selection coefficients, such as s = 0.0407, are sufficient to maintain the observed LD for the most common haplotypes, albeit leading to an unstable equilibrium. Simulations further reveal that the introduction of overdominance stabilizes the system, enabling the long‐term persistence of this complex inversion polymorphism across various frequency scenarios. The stability of this system appears to hinge on a delicate balance between LD, recombination rates, and selective pressures, with overdominance playing a critical role. Our findings highlight the significance of epistatic interactions and selective pressures in shaping evolutionary pathways in natural populations and offer a compelling example of natural selection acting on a complex inversion polymorphism, providing valuable insights into the evolutionary dynamics governing inversion systems.

Biogeography confounds the signal of cospeciation in Batesian mimicry.

Since the inception of the field of evolution, mimicry has yielded insights into foundational evolutionary processes, including adaptive peak shifts, speciation, and the emergence and maintenance of phenotypic polymorphisms.1,2,3 In recent years, the coevolutionary processes generating mimicry have gained increasing attention from researchers. Despite significant advances in understanding Batesian and Müllerian mimicry in Lepidopteran systems, few other mimetic systems have received similar detailed research. Here, we present a Batesian mimicry complex involving flightless, armored Pachyrhynchus weevils and their winged Doliops longhorn beetle mimics and examine their coevolutionary patterns within the Philippine archipelagos. Pachyrhynchus weevils are primarily found in the Philippines, where distinct species radiations have occurred on different islands, each with unique color patterns serving as a warning to predators. This defensive trait and mimicry between unrelated species were first described by Wallace in 1889. Notably, the distantly related longhorn beetle Doliops, despite being soft-bodied and ostensibly palatable, mimics the heavily armored, flightless Pachyrhynchus. To address mimicry in this system, we reconstructed the phylogeny of Doliops using a probe set consisting of 38,000 ultraconserved elements. Our study examines the following questions central to understanding the Pachyrhynchus-Doliops mimicry system: (1) to what extent are coevolutionary interactions conserved (i.e., lineage-constrained) and (2) are the codiversification patterns primarily driven by biotic or abiotic factors?4,5,6 To assess color mimicry and cospeciation, we examined the evolution of nanostructure-based warning colors and the effect of island biogeography on cospeciation. Our findings demonstrate the beetle's ability to repeatedly evolve multiple solutions to similar evolutionary challenges, evolving similar color patterns using different types of photonic crystals with varying degrees of order. We revealed that the observed pattern of cospeciation is driven mainly by abiotic factors from their biogeographic history. Unlike the patterns of coevolution seen between angiosperms and insect lineages,7 most ecological interactions do not persist longer than a few million years, leading to patterns of modularity rather than ecological nestedness.4,6,7.

Territorial status is explained by covariation between boldness, exploration, and thermal preference in a colour polymorphic lizard.

Colour polymorphic species often exhibit variation in morphology, physiology, and behaviour among morphs. In particular, dominance status may be signalled by the interaction between behaviour and colour morph. Behavioural traits associated with dominance include boldness, exploration, and aggression, which influence access to preferred habitat, territorial defence, and mate acquisition. In ectotherms, the social structure associated with morphs may result in the exploitation of structural niches differing in thermal quality. Hence, social interactions among morphs may generate concordant variation in thermal preference and environmental temperature. However, few studies have assessed thermal preference variation in colour polymorphic species and its covariation with behaviour. Doing so can provide insight into niche specialization and the maintenance of colour polymorphism in populations. Here, we investigated the patterns of covariation in boldness behaviour, exploratory behaviour, and thermal preference in the tree lizard, Urosaurus ornatus. We assessed trait variation between territorial and non-territorial male morphs and between orange and yellow female morphs. Boldness and exploratory behaviour were repeatable in male U. ornatus and bolder individuals were significantly more likely to incur tail loss, a potential consequence of bold behaviour. Territorial male morphs were significantly bolder and more exploratory and preferred higher body temperatures with a narrower T set than non-territorial morphs. Female morphs did not vary in behavioural or thermal traits. This study highlights behavioural mechanisms that underly ecological niche segregation and variable habitat use between morphs in a colour polymorphic species.

Mitonuclear effects on sex ratio persist across generations in interpopulation hybrids.

Eukaryotic energy production requires tight coordination between nuclear and mitochondrial gene products. Because males and females often have different energetic strategies, optimal mitonuclear coordination may be sex-specific. Previous work found evidence for sex-specific mitonuclear effects in the copepod Tigriopus californicus by comparing two parental lines and their reciprocal F1 crosses. However, an alternative hypothesis is that the patterns were driven by the parental source of nuclear alleles. Here we test this alternative hypothesis by extending the same cross to F2 hybrids, who receive both maternal and paternal nuclear alleles from F1 hybrids. Results confirm mitonuclear effects on sex ratio, with distorted ratios persisting from the F1 to F2 generations, despite reduced fitness in F2 hybrids. No sex by cross interactions were found for other phenotypic traits measured. Mitochondrial DNA content was higher in females. Both routine metabolic rate and oxidative DNA damage were lower in F2 hybrids than in parentals. The persistence of sex-specific mitonuclear effects, even in the face of F2 hybrid breakdown, attests to the magnitude of these effects, which contribute to the maintenance of within-population mitochondrial DNA polymorphisms.

Structural genomic variation and behavioral interactions underpin a balanced sexual mimicry polymorphism

How phenotypic diversity originates and persists within populations are classic puzzles in evolutionary biology. While balanced polymorphisms segregate within many species, it remains rare for both the genetic basis and the selective forces to be known, leading to an incomplete understanding of many classes of traits under balancing selection. Here, we uncover the genetic architecture of a balanced sexual mimicry polymorphism and identify behavioral mechanisms that may be involved in its maintenance in the swordtail fish Xiphophorus birchmanni. We find that ∼40% of X.birchmanni males develop a "false gravid spot," a melanic pigmentation pattern that mimics the "pregnancy spot" associated with sexual maturity in female live-bearing fish. Using genome-wide association mapping, we detect a single intergenic region associated with variation in the false gravid spot phenotype, which is upstream of kitlga, a melanophore patterning gene. By performing long-read sequencing within and across populations, we identify complex structural rearrangements between alternate alleles at this locus. The false gravid spot haplotype drives increased allele-specific expression of kitlga, which provides a mechanistic explanation for the increased melanophore abundance that causes the spot. By studying social interactions in the laboratory and in nature, we find that males with the false gravid spot experience less aggression; however, they also receive increased attention from other males and are disdained by females. These behavioral interactions may contribute to the maintenance of this phenotypic polymorphism in natural populations. We speculate that structural variants affecting gene regulation may be an underappreciated driver of balanced polymorphisms across diverse species.

The 'queen of the Andes' (Puya raimondii) is genetically fragile and fragmented: a consequence of long generation time and semelparity?

Understanding how life history shapes genetic diversity is a fundamental issue in evolutionary biology, with important consequences for conservation. However, we still have an incomplete picture of the impact of life history on genome-wide patterns of diversity, especially in long-lived semelparous plants. Puya raimondii is a high-altitude semelparous species from the Andes that flowers at 40-100 years of age. We sequenced the whole genome and estimated the nucleotide diversity of 200 individuals sampled from nine populations. Coalescent-based approaches were then used to infer past population dynamics. Finally, these results were compared with results obtained for the iteroparous species, Puya macrura. The nine populations of P. raimondii were highly divergent, highly inbred, and carried an exceptionally high genetic load. They are genetically depauperate, although, locally in the genome, balancing selection contributed to the maintenance of genetic polymorphism. While both P. raimondii and P. macrura went through a severe bottleneck during the Pleistocene, P. raimondii did not recover from it and continuously declined, while P. macrura managed to bounce back. Our results demonstrate the importance of life history, in particular generation time and reproductive strategy, in affecting population dynamics and genomic variation, and illustrate the genetic fragility of long-lived semelparous plants.

Inversions Can Accumulate Balanced Sexual Antagonism: Evidence from Simulations and Drosophila Experiments.

Chromosomal inversion polymorphisms can be common, but the causes of their persistence are often unclear. We propose a model for the maintenance of inversion polymorphism, which requires that some variants contribute antagonistically to two phenotypes, one of which has negative frequency-dependent fitness. These conditions yield a form of frequency-dependent disruptive selection, favoring two predominant haplotypes segregating alleles that favor opposing antagonistic phenotypes. An inversion associated with one haplotype can reduce the fitness load incurred by generating recombinant offspring, reinforcing its linkage to the haplotype and enabling both haplotypes to accumulate more antagonistic variants than expected otherwise. We develop and apply a forward simulator to examine these dynamics under a tradeoff between survival and male display. These simulations indeed generate inversion-associated haplotypes with opposing sex-specific fitness effects. Antagonism strengthens with time, and can ultimately yield karyotypes at surprisingly predictable frequencies, with striking genotype frequency differences between sexes and between developmental stages. To test whether this model may contribute to well-studied yet enigmatic inversion polymorphisms in Drosophila melanogaster, we track inversion frequencies in laboratory crosses to test whether they influence male reproductive success or survival. We find that two of the four tested inversions show significant evidence for the tradeoff examined, with In(3R)K favoring survival and In(3L)Ok favoring male reproduction. In line with the apparent sex-specific fitness effects implied for both of those inversions, In(3L)Ok was also found to be less costly to the viability and/or longevity of males than females, whereas In(3R)K was more beneficial to female survival. Based on this work, we expect that balancing selection on antagonistically pleiotropic traits may provide a significant and underappreciated contribution to the maintenance of natural inversion polymorphism.

The Emergence of Non-Linear Evolutionary Trade-offs and the Maintenance of Genetic Polymorphisms.

Evolutionary models of quantitative traits often assume trade-offs between beneficial and detrimental traits, requiring modelers to specify a function linking costs to benefits. The choice of trade-off function is often consequential; functions that assume diminishing returns (accelerating costs) typically lead to single equilibrium genotypes, while decelerating costs often lead to evolutionary branching. Despite their importance, we still lack a strong theoretical foundation to base the choice of trade-off function. To address this gap, we explore how trade-off functions can emerge from the genetic architecture of a quantitative trait. We developed a multi-locus model of disease resistance, assuming each locus had random antagonistic pleiotropic effects on resistance and fecundity. We used this model to generate genotype landscapes and explored how additive versus epistatic genetic architectures influenced the shape of the trade-off function. Regardless of epistasis, our model consistently led to accelerating costs. We then used our genotype landscapes to build an evolutionary model of disease resistance. Unlike other models with accelerating costs, our approach often led to genetic polymorphisms at equilibrium. Our results suggest that accelerating costs are a strong null model for evolutionary trade-offs and that the eco-evolutionary conditions required for polymorphism may be more nuanced than previously believed.

Carry-over effects of seasonal migration on reproductive success through breeding site retention in a partially migratory bird.

Understanding the maintenance and dynamics of phenotypic polymorphisms requires unpicking key ecological mechanisms shaping the fitness costs and benefits of expressing alternative phenotypes, generating selection. Seasonal migration versus year-round residence expressed in partially migratory populations represents one common polymorphism that can experience strong selection through differential reproductive success. Yet, key hypothesised pathways that could generate such selection remain to be empirically tested. One hypothesis is that migratory tactics affect subsequent reproductive success through carry-over effects on breeding site retention and resulting breeding dispersal. By remaining in breeding areas all year round, residents could retain their preferred breeding site between years, and consequently have higher reproductive success. Conversely, migrants that escape harsh non-breeding season conditions could return in better condition, with high resource holding potential, and outcompete residents to retain their site. Such effects could further depend on migration timing and vary between years. Yet, such pathways have not been quantified, precluding empirical parameterisation of partial migration theory. We used 4 years of breeding and non-breeding season data from partially migratory European shags (Gulosus aristotelis) to test whether the three most frequent migratory tactics in this population (full resident, early migrant departing soon after breeding, and late migrant departing in late autumn) differed in their breeding site retention; whether site retention predicted reproductive success; and hence whether effects of migratory tactic on reproductive success were explicable through site retention. Overall, residents were much more likely to retain their breeding site between years than both early and late migrants, and site retention was associated with increased reproductive success. Yet, these effects varied somewhat among years: late migrants were always least likely to retain their site but had variable relative reproductive success. Path analyses revealed that effects of migratory tactic on reproductive success were only partly attributable to breeding site retention. These results indicate that multiple mechanisms underlie reproductive selection on migratory tactics, potentially contributing to maintaining behavioural polymorphisms. Yet, the clear associations between migratory tactics and local breeding dispersal reveal that these movements can be strongly interlinked across seasons, shaping overall spatioseasonal dynamics in partially migratory systems.

The role of historical biogeography in shaping colour morph diversity in the common wall lizard.

The maintenance of polymorphisms often depends on multiple selective forces, but less is known on the role of stochastic or historical processes in maintaining variation. The common wall lizard (Podarcis muralis) is a colour polymorphic species in which local colour morph frequencies are thought to be modulated by natural and sexual selection. Here, we used genome-wide single-nucleotide polymorphism data to investigate the relationships between morph composition and population biogeography at a regional scale, by comparing morph composition with patterns of genetic variation of 54 populations sampled across the Pyrenees. We found that genetic divergence was explained by geographic distance but not by environmental features. Differences in morph composition were associated with genetic and environmental differentiation, as well as differences in sex ratio. Thus, variation in colour morph frequencies could have arisen via historical events and/or differences in the permeability to gene flow, possibly shaped by the complex topography and environment. In agreement with this hypothesis, colour morph diversity was positively correlated with genetic diversity and rates of gene flow and inversely correlated with the likelihood of the occurrence of bottlenecks. Concurrently, we did not find conclusive evidence for selection in the two colour loci. As an illustration of these effects, we observed that populations with higher proportions of the rarer yellow and yellow-orange morphs had higher genetic diversity. Our results suggest that processes involving a decay in overall genetic diversity, such as reduced gene flow and/or bottleneck events have an important role in shaping population-specific morph composition via non-selective processes.

Pleiotropic effects of melanin pigmentation: haemoparasite infection intensity but not telomere length is associated with plumage morph in black sparrowhawks.

There is increasing recognition of the potential pleiotropic effects of melanin pigmentation, particularly on immunity, with reports of variation in haemoparasite infection intensity and immune responses between the morphs of colour-polymorphic bird species. In a population of the black sparrowhawk (Accipiter melanoleucus) in western South Africa, light morphs have a higher haemoparasite infection intensity, but no physiological effects of this are apparent. Here, we investigate the possible effects of haemoparasite infection on telomere length in this species and explore whether relative telomere length is associated with either plumage morph or sex. Using quantitative polymerase chain reaction analysis, we confirmed that dark morphs had a lower haemoparasite infection intensity than light morphs. However, we found no differences in telomere length associated with either the haemoparasite infection status or morph in adults, although males have longer telomeres than females. While differences in haemoparasite intensity between morphs are consistent with pleiotropic effects of melanin pigmentation in the black sparrowhawk, we found no evidence that telomere length was associated with haemoparasite infection. Further work is needed to investigate the implications of possible pleiotropic effects of plumage morph and their potential role in the maintenance of colour polymorphism in this species.

Climate heterogeneity, season variation, and sexual dimorphism modulate the association between MHC II diversity and parasite variation in striped hamster.

Parasite-mediated selection is widely believed to play a crucial role in maintaining the diversity of the major histocompatibility complex (MHC) genes, which is thought to be maintained through heterozygote advantage, rare-allele advantage, and fluctuating selection. However, the relationship between parasite pressure and MHC diversity has yielded inconsistent findings. These inconsistencies may arise from the influence of environmental factors and individual variations in traits on host-parasite interactions. To address these issues, our study extensively investigated populations of striped hamsters inhabiting regions characterized by environmental heterogeneity. The primary objective was to examine the universality of parasite-mediated selection mechanisms. Our observations revealed the presence of multiple parasite infections, accompanied by spatial and temporal variations in parasite communities and infection patterns among individual hamsters. Specifically, the temperature was found to influence all four parasite indices, while the presence of gamasid mites and parasite richness decreased with increasing precipitation. We also noted significant seasonal variation in parasite dynamics. Moreover, a significant sexual dimorphism was observed with males exhibiting a considerably higher parasite burden compared to their female counterparts. Lastly, we identified the maintenance of MHC polymorphism in striped hamsters as being driven by the heterozygote advantage and fluctuating selection mechanisms. This study underscores the significance of ecological processes in comprehending host-parasite systems and highlights the necessity of considering environmental factors and individual traits when elucidating the mechanisms underlying MHC diversity mediated by parasites.

A cosmopolitan inversion facilitates seasonal adaptation in overwintering Drosophila.

Fluctuations in the strength and direction of natural selection through time are a ubiquitous feature of life on Earth. One evolutionary outcome of such fluctuations is adaptive tracking, wherein populations rapidly adapt from standing genetic variation. In certain circumstances, adaptive tracking can lead to the long-term maintenance of functional polymorphism despite allele frequency change due to selection. Although adaptive tracking is likely a common process, we still have a limited understanding of aspects of its genetic architecture and its strength relative to other evolutionary forces such as drift. Drosophila melanogaster living in temperate regions evolve to track seasonal fluctuations and are an excellent system to tackle these gaps in knowledge. By sequencing orchard populations collected across multiple years, we characterized the genomic signal of seasonal demography and identified that the cosmopolitan inversion In(2L)t facilitates seasonal adaptive tracking and shows molecular footprints of selection. A meta-analysis of phenotypic studies shows that seasonal loci within In(2L)t are associated with behavior, life history, physiology, and morphological traits. We identify candidate loci and experimentally link them to phenotype. Our work contributes to our general understanding of fluctuating selection and highlights the evolutionary outcome and dynamics of contemporary selection on inversions.

Experimental evidence of mate choice as the driving mechanism behind negative assortative mating for shell colour in a marine snail

Abstract How colour polymorphisms are maintained in natural populations constitutes a key subject of study for evolutionary ecologists. One of the mechanisms that has been put forward to explain this phenomenon is negative frequency-dependent selection, which can be caused by different mechanisms. Among these mechanisms is mate choice, or the selection as a potential mate of the rare or dissimilar colour morph by the choosy sex of a given species. In the context of colour polymorphism, mate choice predicts the occurrence of a negative assortative mating pattern, whereby different colour morphs mate more often than expected by chance alone. However, negative assortative mating can also result from other mechanisms, making it, at best, a fallible indicator of a mate choice-driven negative frequency-dependent selection. For this reason, experimental evidence explicitly connecting such a mating pattern to mate choice is necessary to claim that a particular colour polymorphism is being maintained by negative frequency-dependent selection. Here we present experimental evidence backing this relationship for the colour polymorphic marine gastropod Littorina saxatilis from the Ría de Vigo (NW Iberian Peninsula), although how specifically these organisms choose their mates remains unclear. This calls for further experimental efforts to clarify this issue and its relationship to the maintenance of colour polymorphism in L. saxatilis.

Overcoming the congenitally disadvantageous mutation through adaptation to environmental UV exposure in land snails.

Congenital fitness-disadvantageous mutations are not maintained in the population; they are purged from the population through processes such as purifying selection. However, these mutations could persist in the population as polymorphisms when it is advantageous for the individuals carrying them in adapting to a specific external environment. We tested this hypothesis using the dimorphic land snail Euhadra peliomphala simodae in Japan; these snails have dark or bright coloured shells. The survival rate of dark snails at hatching was lower than that of the bright ones, as observed in the F1 progenies produced through crossing. Dark snails have a congenital fitness-disadvantageous mutation; however, they also have protection against ultraviolet radiation. They have a higher survival rate than the bright snails in a UV environment, as observed using the UV exposure experiments and UV transmittance measurements. This is a good example of a congenitally disadvantageous mutation that is advantageous for adapting to the external environment. These results explain the maintenance of polymorphism and highlight the genotypic and phenotypic diversity in the wild population.

Colour-specific diet specialization is associated with differences in owlet weight in a polymorphic owl: influence of the trophic quality variation.

The niche divergence hypothesis proposes that the evolution and maintenance of colour polymorphism is based on a mechanism of disruptive selection. In a trophic context, the hypothesis predicts that individuals differing in colour vary in their trophic niche, either because they differ in foraging efficiency or feed in different habitats. A major evolutionary conundrum is how these expectations are affected by variation in trophic quality. Using an owl species with colour plumage polymorphism, the Eurasian scops owl Otus scops, we examined diet and habitat segregation during reproduction in relation to plumage colouration and trophic quality. Intensive sampling revealed that trophic quality for scops owls (i.e. abundance of grasshoppers and locusts) varied more among territories than between years, but scops owls did not segregate among territories of different quality by their colouration. However, we found that sex, plumage colouration and territory differences in trophic quality explained differences in the degree of dietary specialization. Brownish males delivered a higher diversity of prey to the nest than greyish ones in high trophic quality territories. We also found that the more diverse the diet provided by males, the heavier the owlets at fledging. Our study provides evidence for a different sensitivity to trophic quality of the colour morphs with potential fitness consequences in scops owls. We highlight the importance of studying the mechanisms leading to the persistence of colour polymorphism in patchy environments, since segregation may pass otherwise unnoticed if only habitats or years with similar conditions are considered.

Nutrition regulates sex expression in a gender diphasy plant, Lilium concolor var. megalanthum.

The evolution and maintenance of plant polymorphism have always received much attention. Gender diphasy is a rare sexual system. Plant individuals with gender diphasy can adjust the resource allocation of different functional organs according to the changes of environmental conditions to regulate the sex expression of individuals, and the sex expression can be converted between years. However, our understanding of sex expression in plants is still very insufficient. In this study, we explored whether the perennial plant Lilium concolor var. megalanthum has a gender diphasy system and whether environmental resource availability affects its resource allocation and sex expression. By collecting the bulbs of two sexual phenotypes (male and hermaphrodite) in the field and simulating the application of different levels of nutrients under the same habitat conditions, the growth and reproduction indexes and sex expression of plants in two years (2021 and 2022) were measured to evaluate the resource allocation strategy and sex expression pattern of Lilium concolor var. megalanthum. We found that the sex expression of Lilium concolor var. megalanthum was variable in continuous years. Under limited resources, Lilium concolor var. megalanthum increases the biomass resources of the leaves and has a longer flowering period. Resource availability regulates the growth, reproduction and sex expression of Lilium concolor var. megalanthum. Bulb size is the main factor affecting its growth, reproduction and sex expression after accumulating sufficient resources. This study confirms that Lilium concolor var. megalanthum has gender diphasy system. There is a strong trade-off between growth and reproduction under limited resources. Nutrient levels can regulate the reproduction and sex expression process of Lilium concolor var. megalanthum. With the growth of Lilium concolor var. megalanthum in consecutive years, the size dependence of bulbs may be the decisive factor in its sex expression.