Assessing ecological and physiological costs of melanism in North American Papilio glaucus females: two decades of dark morph frequency declines

Polymorphisms for melanic form of insects may provide various selective advantages. However, melanic alleles may have significant/subtle pleiotrophic “costs.” Several potential pleiotrophic effects of the W (=Y)‐linked melanism gene in Papilio glaucus L. (Lepidoptera) showed no costs for melanic versus yellow in adult size, oviposition preferences, fecundity, egg viability, larval survival/growth rates, cold stress tolerance, or postdiapause emergence times. Sexual selection (males choosing yellow rather than mimetic dark females) had been suggested to provide a balanced polymorphism in P. glaucus, but spermatophore counts in wild females and direct field tethering studies of size‐matched pairs of virgin females (dark and yellow), show that male preferences are random or frequency‐dependent from Florida to Michigan, providing no yellow counter‐advantages. Recent frequency declines of dark (melanic/mimetic) females in P. glaucus populations are shown in several major populations from Florida (27.3°N latitude) to Ohio (38.5° N). Summer temperatures have increased significantly at all these locations during this time (1999–2018), but whether dark morphs may be more vulnerable (in any stage) to such climate warming remains to be determined. Additional potential reasons for the frequency declines in mimetic females are discussed: (i) genetic introgression of Z‐linked melanism suppressor genes from P. canadensis (R & J) and the hybrid species, P. appalachiensis (Pavulaan & Wright), (ii) differential developmental incompatibilities, or Haldane effects, known to occur in hybrids, (iii) selection against intermediately melanic (“dusty”) females (with the W‐linked melanic gene, b+) which higher temperatures can cause.     

Amplified fragment length polymorphism analysis of Mythimna separata (Lepidoptera: Noctuidae) geographic and melanic laboratory populations in China

Genetic diversity within and among three wild-type natural populations and one melanic laboratory population of Mythimna separata (Walker) (Lepidoptera: Noctuidae) were evaluated using amplified fragment length polymorphism (AFLP) analysis. Although extensive genetic diversity occurs among individuals from different geographic populations (P = 54.5%, h = 0.209, I = 0.305), the majority of the genetic diversity is within populations and not between populations (G(ST) = 0.172), indicating high gene flow (N(M) = 2.403) and suggesting that M. separata in northern China are a part of a single large metapopulation. Genetic diversity in the natural populations was significantly higher than that in the melanic laboratory population (with P = 43.4% versus P = 25.9%, h = 0.173 versus h = 0.086, and I = 0.251 versus I = 0.127), suggesting that the melanic laboratory population is narrowly genetic-based and genetically uniform. Genetic similarities based on AFLP data were calculated, and cluster analysis was preformed to graphically display groupings between individuals and populations. Individuals from the same region were not grouped together in cluster analysis of three natural populations, whereas melanic individuals from laboratory population were grouped together very well. Four subpopulations were clustered into two broad groups. Melanic laboratory population became a single group, which had apparent differentiation from the other group in which three natural subpopulations were included. These results indicated that although high genetic variability existed among the individuals of natural populations, there was little genetic differentiation among three geographic populations that could be explained by the effects of the long distance migration of the oriental armyworm in China enhanced the level of gene flow. Influences of migration on the genetic polymorphism and differentiations that make a significant contribution to evolution in this insect are reviewed.     

A study of nonrandom mating in a British population of the two-spot ladybird with a high frequency of the melanic morph

In some studies of the two-spot ladybird (Adalia bipunctata), melanic males have been found in excess over the typical morph in matings. Data suggest that a genetic female mating preference is responsible. The mating advantage of melanic males may be important in maintaining a polymorphism between melanic and typical ladybirds in many populations in the United Kingdom (U.K.). It has been reported that preference frequency varies linearly with melanic frequency throughout most of the U.K. One particular population ofAdalia bipunctata near Aberdare, South Wales, is noted for its high frequency of melanic individuals. It has been suggested that local environmental factors account for the high melanic frequency in this population. It is also possible, however, that a female mating preference may be at least partly responsible for the high frequency of melanics (as has been proposed for the rest of the U.K.). In this study, experiments have been performed to determine the level of female mating preference in the Aberdare population. No evidence was found for any mating advantage to melanic males. There was inconsistent and unexpected evidence that melanic females were overrepresented in matings, but the cause for this was unclear. Female mating preference does not appear, therefore, to be responsible for the high melanic frequency in the population ofAdalia bipunctata near Aberdare. There is not a simple association between mating preference and melanic frequency in U. K. populations of the two-spot ladybird.     

The genetic underpinnings of population cyclicity: establishing expectations for the genetic anatomy of cycling populations

Despite extensive research into the mechanisms underlying population cyclicity, we have little understanding of the impacts of numerical fluctuations on the genetic variation of cycling populations. Thus, the potential implications of natural and anthropogenically‐driven variation in population cycle dynamics on the diversity and evolutionary potential of cyclic populations is unclear. Here, we use Canada lynx Lynx canadensis matrix population models, set up in a linear stepping‐stone, to generate demographic replicates of biologically realistic cycling populations. Overall, increasing cycle amplitude predictably reduced genetic diversity and increased genetic differentiation, with cyclic effects increased by population synchrony. Modest dispersal rates (1–3% of the population) between high and low amplitude cyclic populations did not diminish these effects suggesting that spatial variation in cyclic amplitude should be reflected in patterns of genetic diversity and differentiation at these rates. At high dispersal rates (6%) groups containing only high amplitude cyclic populations had higher diversity and lower differentiation than those mixed with low amplitude cyclic populations. Negative density‐dependent dispersal did not impact genetic diversity, but did homogenize populations by reducing differentiation and patterns of isolation by distance. Surprisingly, temporal changes in diversity and differentiation throughout a cycle were not always consistent with population size. In particular, negative density‐dependent dispersal simultaneously decreased differences in genetic diversity while increasing differences in genetic differentiation between numerical peaks and nadirs. Combined, our findings suggest demographic changes at fine temporal scales can impact genetic variation of interacting populations and provide testable predictions relating population cyclicty to genetic variation. Further, our results suggest that including realistic demographic and dispersal parameters in population genetic models and using information from temporal changes in genetic variation could help to discern complex demographic scenarios and illuminate population dynamics at fine temporal scales.     

Life-history changes that accompany the transition from sexual to parthenogenetic reproduction in Drosophila mercatorum

In spite of the predicted genetic and ecological costs of sex, most natural populations maintain sexual reproduction, even those capable of facultative parthenogenesis. Unfertilized eggs from natural populations of Drosophila mercatorum occasionally develop into viable adults, but obligately parthenogenetic populations are unknown in this species. To evaluate the microevolutionary forces that both favor and constrain the evolution of parthenogenesis in D. mercatorum, we have measured parthenogenetic rates across a natural, sexually reproducing population and characterized the life-history changes that accompany the transition from sexual to parthenogenetic reproduction in laboratory strains. A highly significant difference in parthenogenetic rate was found between two populations in close geographic proximity, with increased rate found with lower population density. Laboratory strains of parthenogenetic females suffered increased mortality and reduced egg viability relative to their virgin counterparts from a sexual strain. Lifetime egg production was similar across all strains, but a shift in peak egg production to an earlier age also occurred. The combination of these life-history traits resulted in a higher net reproductive value for sexual females, but because they also had a longer generation time, intrinsic rate of increase was not as dramatically different from parthenogenetic females. In environments with high early mortality, there may be no fitness disadvantage to parthenogenesis, but the predicted ecological advantage of a twofold increase in intrinsic rate of increase was not realized. These results support the theory of Stalker (1956) that parthenogenesis is favored in environments in which sexual reproduction is difficult or impossible.     

Microevolution in an electronic microcosm

The evolution of microbial populations in simple environments such as chemostats is still not fully understood. The classical interpretation of adaptation involves a process of successive substitution whereby a new dominant genotype arises by mutation from the genotype previously dominant and spreads more or less rapidly through the population until it is nearly fixed. The population is, thus, nearly uniform most of the time. Some observations suggest that the process may be more complicated, but it remains formidably difficult to assemble the phylogeny of an evolving culture in sufficient detail to be sure. We report experiments with an electronic microcosm inhabited by self-replicating computer programs whose phylogeny can be rendered completely transparent. The physiology of these programs is different in many respects from that of organic creatures, but their population biology has many features in common, including a very extensive, if not unbounded, range of variation. Experimental populations evolved through point mutations (many of which were quasi-neutral when they were viable) and through rearrangements that led to a change in genome size and often had large effects on fitness. As a general rule, smaller genomes execute fewer instructions in order to replicate, the rate of replication increases as the number of instructions executed declines, and the rate of replication in pure culture is a good predictor of success in mixture. When cultured with CPU (central processing unit) time as the sole limiting resource, smaller genomes, therefore, evolve as a correlated response to natural selection for faster replication. The genetic basis of adaptation was highly contingent and always differed in replicate experiments. The pattern of evolution depends on mutation rate. At low mutation rates of 0.01 per genome per generation or less, we observed classic periodic selection, with each dominant genotype descending from the previous dominant and rising to a frequency of 0.8 or more. At higher mutation rates of about 0.1 per genome per generation, the most abundant genotypes rarely exceeded a frequency of about 0.4, and rare genotypes present in a few copies comprised a large part of the population. New dominant genotypes did not usually descend directly from previous dominants but, instead, from one of the many rare or moderately abundant genotypes. We suggest that the conventional chemostat paradigm may hold only as a special case at very low mutation rates and that the dynamics and diversity of evolving populations, even in the simplest conditions, may be more complex than is usually recognized. Artificial genetic autoadaptive systems are likely to be useful in constructing theory for situations that lie beyond the boundary of conventional population genetics.     

The molecular basis of melanism and mimicry in a swallowtail butterfly

Melanism in Lepidoptera, either industrial or in mimicry, is one of the most commonly cited examples of natural selection [1] [2]. Despite extensive studies of the frequency and maintenance of melanic genes in insect populations [1] [2], there has been little work on the underlying molecular mechanisms. Nowhere is butterfly melanism more striking than in the Eastern Tiger Swallowtail (Papilio glaucus) of North America [3] [4] [5]. In this species, females can be either yellow (wild type) or black (melanic). The melanic form is a Batesian mimic of the distasteful Pipevine Swallowtail (Battus philenor), which is also black in overall color. Melanism in P. glaucus is controlled by a single Y-linked (female) black gene [6]. Melanic females, therefore, always have melanic daughters. Black melanin replaces the background yellow in melanic females. Here, we show that the key enzyme involved is N-beta-alanyl-dopamine-synthase (BAS), which shunts dopamine from the melanin pathway into the production of the yellow color pigment papiliochrome and also provides products for cuticle sclerotization. In melanic females, this enzyme is suppressed, leading to abnormal melanization of a formerly yellow area, and wing scale maturation is also delayed in the same area. This raises the possibility that either reduced BAS activity itself is preventing scale sclerotization (maturation) or, in contrast, that the delay in scale maturation precludes expression of BAS at the correct stage. Together, these data show how changes in expression of a single gene product could result in multiple wing color phenotypes. The implications for the genetic control of mimicry in other Lepidoptera are discussed.     

Consequences of past and present harvest management in a declining flyway population of common eiders Somateria mollissima

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Street fighters: Bite force,injury rates,and density of urban Australian water dragons (Intellagama lesueurii)

In an increasingly urbanized world it is imperative that we understand how wildlife responds to this novel anthropogenic landscape, both at the individual‐ and population‐level. Urbanization generally reduces biodiversity, but can also favour particular species and increase their abundance relative to wild populations. When population density increases, so too does the frequency and cost of social interactions. We studied Australian water dragons (Intellagama lesueurii), a species common in urban, semi‐natural and natural areas, to firstly test the prediction that urban populations occur at higher densities, and then determine the consequences of urbanization for combat rates (quantified using wounding) and bite force. We established that urban populations are denser than ones from semi‐natural and natural habitats. We also recorded significantly more wounds in females from urban populations than females from both natural and semi‐natural populations. Urban males also had significantly higher incidence of wounding than males from natural populations. We did not find a difference in male or female bite force among any populations across the urban‐natural gradient. Overall, we found evidence that urbanization results in a higher population density of water dragons and more frequent conspecific combat, but this was not associated with an increase in bite force. These finding suggests that there may be a physiological cost to living in urban habitats related to increased contest rates and wounding.     

Frequency‐dependent selection for rare genotypes promotes genetic diversity of a tropical palm

Negative frequency‐dependent selection among species is a key driver of community diversity in natural systems, but the degree to which negative frequency‐dependent selection shapes patterns of survival and genetic diversity within species is poorly understood. In a 5‐year field experiment, we show that seedlings of a tropical palm with rare genotypes had a pronounced survival advantage over seedlings with common genotypes, with effect sizes comparable to that of light availability. This ‘rare genotype advantage’ led to an increase in population‐wide genetic diversity among seedlings compared to null expectations, as predicted by negative frequency‐dependent selection, and increased reproductive success in adult trees with rare genotypes. These results suggest that within‐species negative frequency‐dependent selection of genotypes can shape genetic variation on ecologically relevant timescales in natural systems and may be a key, overlooked source of non‐random mortality for tropical plants.     

The maintenance of nucleocytoplasmic polymorphism in a metapopulation: the case of gynodioecy

In gynodioecious species, gender is generally determined by epistatic interactions between cytoplasmic and nuclear loci. However, theoretical studies suggest that, for a joint polymorphism at both cytoplasmic and nuclear loci to be maintained in a panmictic population, selection must act differently on the various genotypes that determine the same gender. Here we show that, in a metapopulation with local extinction and restricted gene flow, nucleocytoplasmic polymorphism can be maintained without these differences. We use deterministic simulations. We assume that gene flow occurred only at recolonization. Founder effects create genetic variance between populations in the metapopulation, and local population growth is faster when the local frequency of females is high. Group selection phenomena are involved in the maintenance of the joint polymorphism in the metapopulation. The frequency of females in the metapopulation at equilibrium is higher than in a panmictic population with the same genetic system. However, these conclusions hold only if nuclear alleles restoring male fertility are dominant.     

The maintenance of mitochondrial genetic variation by negative frequency‐dependent selection

Mitochondrial genes generally show high levels of standing genetic variation, which is puzzling given the accumulating evidence for phenotypic effects of mitochondrial genetic variation. Negative frequency‐dependent selection, where the relative fitness of a genotype is inversely related to its frequency in a population, provides a potent and potentially general process that can maintain mitochondrial polymorphism. We assessed the change in mitochondrial haplotype frequencies over 10 generations of experimental evolution in 180 seed beetle populations in the laboratory, where haplotypes competed for propagation to subsequent generations. We found that haplotypes consistently increased in frequency when they were initially rare and decreased in frequency when initially common. Our results have important implications for the use of mtDNA haplotype frequency data to infer population level processes and they revive the general hypothesis that negative frequency‐dependent selection, presumably caused by habitat heterogeneity, may commonly promote polymorphism in ecologically relevant life history genes.     

Predation and the persistence of melanic male mosquitofish (Gambusia holbrooki)

The empirical reasons for the persistent rarity of a genotype are typically complex and tedious to identify, particularly in nature. Yet rare morphs occur in a substantial fraction of phenotypic polymorphisms. A colour polymorphism has persisted for decades in the eastern mosquitofish, yet why this is so remains obscure. Here, I report the results of (1) intensive sampling at 45 natural sites to obtain the frequency distribution of the melanic (black) mosquitofish morph in Florida, (2) predation trials, conducted independently in mesocosms, with three different predatory species and (3) two mark-recapture studies, conducted in nature. This work (1) documents the rarity of melanic mosquitofish in nature, (2) demonstrates that melanic males experience a selective advantage over silver males in the presence of predators, (3) indicates no difference in the colour morphs' survival at a natural site essentially devoid of these predators, although suggesting a higher rate of recapture for melanic males at a site rife with predators. Overall, selective predation appears to contribute to the persistence of the melanic morph, despite its rarity in nature.     

Variation in and correlation between intrinsic rate of increase and carrying capacity

Intrinsic population growth rate and density dependence are fundamental components of population dynamics. Theory suggests that variation in and correlations between these parameters among patches within a population can influence overall population size, but data on the degree of variation and correlation are rare. Replicate populations of a specialist aphid (Chaetosiphon fragaefolii) were followed on 11 genotypes of host plant (Fragaria chiloensis) in the greenhouse. Population models fit to these census data provide estimates of intrinsic growth rate and carrying capacity for aphid populations on each plant genotype. Growth rate and carrying capacity varied substantially among plant genotypes, and these two parameters were not significantly correlated. These results support the existence of spatial variation in population dynamic parameters; data on frequency distributions and correlations of these parameters in natural populations are needed for evaluation of the importance of variation in growth rate and density dependence for population dynamics in the field.     

Dynamics of a Lactate Dehydrogenase Polymorphism in the Wood Louse PORCELLIO SCABER Latr.: Evidence for Partial Assortative Mating and Heterosis in Natural Populations

Electrophoretic separation of lactate dehydrogenase (LDH) of Porcellio scaber from 14 natural populations in California, and one each in Oregon, Delaware and Massachusetts, indicates a biallelic polymorphism. Phenotypes are recovered from laboratory matings of virgin females in frequencies agreeing with simple Mendelian inheritance, and the frequency distributions of phenotypes in natural populations are typically in agreement with the appropriate Hardy-Weinberg distributions for these same populations. The same allele predominates in all natural populations examined. Temporal stability within populations suggests that the polymorphism is at, or near, equilibrium. The spatial distribution of allele frequencies, however, is apparently mosaic. Abrupt discontinuities in gene frequency over short distances (50 m to 1 km) suggest that interpopulation migration is insufficient to swamp local differences in gene frequency. Analysis of the transmission dynamics of the polymorphism in natural populations using mother-offspring genotype comparisons suggests that the allelic frequencies of transmitted male gametes are not independent of female genotype. Specifically, the observed mating scheme in natural populations appears to be partially assortative. Comparisons of progeny genotype distributions with yearling (or adult) genotype distributions from the same populations indicate a superior post-partum viability of heterozygous individuals relative to homozygotes. The distortion of progeny genotypic distributions created by assortment is thus apparently counteracted by subsequent heterosis.     

Low RAPD Variation and Female-Biased Sex Ratio Indicate Genetic Drift in Small Populations of the Dioecious Conifer Taxus Baccata in Switzerland

Small populations are prone to genetic drift as a consequence of random sampling effects. We investigated whether we could detect such random sampling effects in the English yew (Taxus baccata), a dioecious conifer species occurring in scattered populations in Switzerland. Seven pairs of small and large populations were analyzed using random amplified polymorphic DNA (RAPD) marker bands from 20 individuals per population. Several genetic parameters (mean marker band frequency deviation, molecular variance, population differentiation) indicated that small populations experienced genetic drift. These genetic differences between small and large populations of yew were paralleled by an increased sex ratio bias towards a higher number of females in the small populations. Our findings support earlier assumptions that the Swiss occurrences of yew may be described as metapopulation dynamics, characterized by local colonization and extinction events leading to the observed genetic drift.     

Genetic quality of individuals impacts population dynamics

Ample evidence exists that an increase in the inbreeding level of a population reduces the value of fitness components such as fecundity and survival. It does not follow, however, that these decreases in the components of fitness impact population dynamics in a way that increases extinction risk, because virtually all species produce far more offspring than can actually survive. We analyzed the effects of the genetic quality (mean fitness) of individuals on the population growth rate of seven natural populations in each of two species of wolf spider in the genus Rabidosa , statistically controlling for environmental factors. We show that populations of different sizes, and different inbreeding levels, differ in population dynamics for both species. Differences in population growth rates are especially pronounced during stressful environmental conditions (low food availability) and the stressful environment affects smaller populations (<500 individuals) disproportionately. Thus, even in an invertebrate with an extremely high potential growth rate and strong density-dependent mortality rates, genetic factors contribute directly to population dynamics and, therefore, to extinction risk. This is only the second study to demonstrate an impact of the genetic quality of individual genotypes on population dynamics in a wild population and the first to document strong inbreeding–environment interactions for fitness among populations. Endangered species typically exist at sizes of a few hundred individuals and human activities degrade habitats making them innately more stressful (e.g. global climate change). Therefore, the interaction between genetic factors and environmental stress has important implications for efforts aimed at conserving the Earth's biodiversity.     

The costs of colour: plasticity of melanin pigmentation in an outbreaking polymorphic forest moth

Outside the context of industrial melanism, little is known about the physiological and ecological importance of genetic melanic polymorphisms in moths. Melanin pigments are synthesized from amino acid precursors and should therefore be costly to produce in nitrogen‐limited insects. A genetic melanic polymorphism is present in adult Malacosoma disstria Hübner (Lepidoptera: Lasiocampidae), a widespread forest moth with outbreaking population dynamics. We test the hypotheses that melanin‐based colouration is physiologically costly in M. disstria, that expression of melanin‐based colouration is a plastic trait which varies with population density and nutrition, and that the genetically based melanic phenotype is disadvantaged under nutritionally poor conditions. Two experiments were used to test these hypotheses. A field study compared pigmentation and phenotypic frequencies in moths collected from high‐ and low‐density populations. A laboratory experiment investigated the effects of larval nitrogen availability on adult pigmentation and phenotypic frequencies. High population density and nitrogen limitation reduced pigmentation and size of all moths, but phenotypic frequencies were not affected in either experiment. The effects of diet on both pigmentation and size were stronger for melanic moths than for typical moths. Our results show that adult melanism in M. disstria is physiologically costly, that colour expression is plastic despite its genetic component, and that the melanic phenotype may be disadvantaged under poor conditions but favoured under good conditions. We suggest that temporal variation in selection and trait plasticity help maintain polymorphism stability.     

Mating system contributes only slightly to female maintenance in gynodioecious Geranium maculatum (Geraniaceae)

Gynodioecy, the co-occurrence of female and hermaphroditic individuals within a population, is an important intermediate in the evolution of separate sexes. The first step, female maintenance, requires females to have higher seed fitness compared with hermaphrodites. A common mechanism thought to increase relative female fitness is inbreeding depression avoidance, the magnitude of which depends on hermaphroditic selfing rates and the strength of inbreeding depression. Less well studied is the effect of biparental inbreeding on female fitness. Biparental inbreeding can affect relative female fitness only if its consequence or frequency differs between sexes, which could occur if sex structure and genetic structure both occur within populations. To determine whether inbreeding avoidance and/or biparental inbreeding can account for female persistence in Geranium maculatum, we measured selfing and biparental inbreeding rates in four populations and the spatial genetic structure in six populations. Selfing rates of hermaphrodites were low and did not differ significantly from zero in any population, leading to females gaining at most a 1–14% increase in seed fitness from inbreeding avoidance. Additionally, although significant spatial genetic structure was found in all populations, biparental inbreeding rates were low and only differed between sexes in one population, thereby having little influence on female fitness. A review of the literature revealed few sexual differences in biparental inbreeding among other gynodioecious species. Our results show that mating system differences may not fully account for female maintenance in this species, suggesting other mechanisms may be involved.     

Simulating the maintenance of a rare fish morph experiencing negative frequency dependent selection

Empirical work assessing the maintenance of rare genotypes in natural populations is difficult over very long time scales. Skirting this problematic issue is possible with theory and simulations. Major theoretical constructs, including mutation-selection balance and balancing selection, explain the theoretical maintenance of rare genotypes, and the occurrence of multiple, rare genotypes over time. Additionally, numerical simulations are valuable tools for assessing evolving biological systems because they allow for monitoring systems over long time scales, as well as for controlling model parameters, thus contributing to the exploration of system dynamics that cannot be assessed in nature. Here we employed numerical simulations to explore the importance of several biological factors that contribute to the maintenance of a fish color-pattern polymorphism. We present a numerical model of a two-morph fish polymorphism that allowed us to test the sensitivity of the rare morph's persistence and the population's stability to multiple parameters. Our simulations ran over 10,000 years (where one year is approximately one generation) and demonstrated the maintenance of a stable polymorphism with a rare morph which persisted at a frequency of ∼10⁻², which is in-fact the frequency of the rare, mottled black mosquitofish morph in natural populations. This pigmentation polymorphism is stable, independent of changes in population size, but can be destabilized with very high predation when coupled with very low birth rates. Employing models with empirical fitness estimates is a valuable tool for monitoring rare vertebrate morphs in nature, however few studies exist that have accomplished this task. Our approach can be adapted for modeling rare morphs (particularly in additional live-bearing fishes like sailfin mollies) that also harbor rare, pigmentation morphs within large populations.