The evolution of exploitative specificity can be influenced by environmental variability in space and time and the intensity of trade-offs. Coevolution, the process of reciprocal adaptation in two or more species, can produce variability in host exploitation and as such potentially drive patterns in host and parasite specificity. We employed the bacterium Pseudomonas fluorescens SBW25 and its DNA phage Φ2 to investigate the role of coevolution in the evolution of phage infectivity range and its relation with phage growth rate. At the phage population level, coevolution led to the evolution of broader infectivity range, but without an associated decrease in phage growth rate relative to the ancestor, whereas phage evolution in the absence of bacterial evolution led to an increased growth rate but no increase in infectivity range. In contrast, both selection regimes led to phage adaptation (in terms of growth rates) to their respective bacterial hosts. At the level of individual phage genotypes, coevolution resulted in within-population diversification in generalist and specialist infectivity range types. This pattern was consistent with a multilocus gene-for-gene interaction, further confirmed by an observed cost of broad infectivity range for individual phage. Moreover, coevolution led to the emergence of bacterial genotype by phage genotype interactions in the reduction of bacterial growth rate by phage. Our study demonstrates that the strong reciprocal selective pressures underlying the process of coevolution lead to the emergence and coexistence of different strategies within populations and to specialization between selective environments.

Divergence between populations adapting to different environments may be facilitated when the populations differ in their sexual traits. We tested whether colonizing a novel environment may, through phenotypic plasticity, change sexual traits in a way that could alter the dynamics of sexual selection. This hypothesis has two components: changes in mean phenotypes across environments, and changes in the genetic background of the phenotypes that are produced—or genotype × environment interaction (G × E). We simulated colonization of a novel environment and tested its effect on the mating signals of a member of the Enchenopa binotata species complex of treehoppers (Hemiptera: Membracidae), a clade that has diverged in a process involving host plant shifts and signal diversification. We found substantial genetic variation and G × E in most signal traits measured, with little or no change in mean signal phenotypes. We suggest that the expression of extant genetic variation across old and novel environments can initiate signal divergence.

Predicting the host range for herbivores has been a major aim of research into plant–herbivore interactions and an important model system for understanding the evolution of feeding specialization. Among many terrestrial insects, host range is strongly affected by herbivore phylogeny and long historical associations between particular herbivore and plant taxa. For small herbivores in marine environments, it is known that the evolution of host use is sculpted by several ecological factors (e.g., food quality, value as a refuge from predators, and abiotic forces), but the potential for phylogenetic constraints on host use remains largely unexplored. Here, we analyze reports of host use of herbivorous amphipods from the family Ampithoidae (102 amphipod species from 12 genera) to test the hypotheses that host breadth and composition vary among herbivore lineages, and to quantify the extent to which nonpolar secondary metabolites mediate these patterns. The family as a whole, and most individual species, are found on a wide variety of macroalgae and seagrasses. Despite this polyphagous host use, amphipod genera consistently differed in host range and composition. As an example, the genus Peramphithoe rarely use available macrophytes in the order Dictyotales (e.g., Dictyota) and as a consequence, display a more restricted host range than do other genera (e.g., Ampithoe, Cymadusa, or Exampithoe). The strong phylogenetic effect on host use was independent of the uneven distribution of host taxa among geographic regions. Algae that produced nonpolar secondary metabolites were colonized by higher numbers of amphipod species relative to chemically poor genera, consistent with the notion that secondary metabolites do not provide algae an escape from amphipod herbivory. In contrast to patterns described for some groups of phytophagous insects, marine amphipods that use chemically rich algae tended to have broader, not narrower, host ranges. This result suggests that an evolutionary advantage to metabolite tolerance in marine amphipods may be that it increases the availability of appropriate algal hosts (i.e., enlarges the resource base).

Gene exchange between individuals can lead to profound evolutionary effects at both the genomic and population levels. These effects have sparked widespread interest in examining the specific adaptive benefits of recombination. Although this work has primarily focused on the benefits of sex in eukaryotes, it is assumed that similar benefits of genetic exchange apply across eukaryotes and prokaryotes. Here we report a direct test of this assumption using the naturally transformable human gastric pathogen Helicobacter pylori as a model organism. We show that genetic exchange accelerates adaptation to a novel laboratory environment within bacterial populations and that a general adaptive advantage exists for naturally transformable strains when transfer occurs among conspecific backgrounds. This finding demonstrates that there are generalized benefits to adaptation in both eukaryotes and prokaryotes even though the underlying processes are mechanistically different.

House mice offer a powerful system for dissecting the genetic basis of phenotypes that isolate species in the early stages of speciation. We used a series of reciprocal crosses between wild-derived strains of Mus musculus and M. domesticus to examine F₁ hybrid male sterility, one of the primary phenotypes thought to isolate these species. We report four main results. First, we found significantly smaller testes and fewer sperm in hybrid male progeny of most crosses. Second, in some crosses hybrid male sterility was asymmetric and depended on the species origin of the X chromosome. These observations confirm and extend previous findings, underscoring the central role that the M. musculus X chromosome plays in reproductive isolation. Third, comparisons among reciprocal crosses revealed polymorphism at one or more hybrid incompatibilities within M. musculus. Fourth, the spermatogenic phenotype of this polymorphic interaction appears distinct from previously described hybrid incompatibilities between these species. These data build on previous studies of speciation in house mice and show that the genetic basis of hybrid male sterility is fairly complex, even at this early stage of divergence.

Clinally varying traits in Drosophila melanogaster provide good opportunities for elucidating the genetic basis of adaptation. Resistance to ethanol, a natural component of D. melanogaster's breeding sites, increases with latitude on multiple continents, indicating that the trait is under selection. Although the well-studied Alcohol dehydrogenase (Adh) polymorphism makes a contribution to the clines, it accounts for only a small proportion of the phenotypic variation. We describe an amino acid replacement polymorphism in Aldehyde dehydrogenase (Aldh), the gene encoding the second enzyme in the ethanol degradation pathway, that shows hallmarks of also contributing to the clines. The derived Aldh allele, like the Adh-Fast allele, increases in frequency in laboratory populations selected for ethanol resistance, and increases in frequency with latitude in wild populations. Moreover, strains with the derived allele have significantly higher ALDH enzyme activity with acetaldehyde (the breakdown product of ethanol) as a substrate than strains with the ancestral allele. As is the case with the Adh-Fast allele, chromosomes with the derived Aldh allele show markedly reduced molecular variation in the vicinity of the replacement polymorphism compared to those with the ancestral allele, suggesting a single, relatively recent origin. Nonetheless, the Aldh polymorphism differs from the Adh polymorphism in that the ethanol-associated allele remains in relatively low frequency in most populations. We present evidence that this is likely to be the result of a trade-off in catalytic activity, with the advantage of the derived allele in acetaldehyde detoxification being offset by a disadvantage in detoxification of other aldehydes.

The distribution of effect sizes of genes underlying adaptation is unknown (Orr 2005). Are suites of traits that diverged under natural selection controlled by a few pleiotropic genes of large effect (major genes model), by many independently acting genes of small effect (infinitesimal model), or by a combination, with frequency inversely related to effect size (geometric model)? To address this we carried out a quantitative trait loci (QTL) study of a suite of 54 position traits describing body shapes of two threespine stickleback species: an ancestral Pacific marine form and a highly derived benthic species inhabiting a geologically young lake. About half of the 26 detected QTL affected just one coordinate and had small net effects, but several genomic regions affected multiple aspects of shape and had large net effects. The distribution of effect sizes followed the gamma distribution, as predicted by the geometric model of adaptation when detection limits are taken into account. The sex-determining chromosome region had the largest effect of any QTL. Ancestral sexual dimorphism was similar to the direction of divergence, and was largely eliminated during freshwater adaptation, suggesting that sex differences may provide variation upon which selection can act. Several shape QTL are linked to Eda, a major gene responsible for reduction of lateral body armor in freshwater. Our results are consistent with predictions of the geometric model of adaptation. Shape evolution in stickleback results from a few genes with large and possibly widespread effects and multiple genes of smaller effect.

Understanding the evolution of sexual ornaments, and particularly that of female sexual ornaments, is an enduring challenge in evolutionary biology. Key to this challenge are establishing the relationship between ornament expression and female reproductive investment, and determining the genetic basis underpinning such relationship. Advances in genomics provide unprecedented opportunities to study the genetic architecture of sexual ornaments in model species. Here, we present a quantitative trait locus (QTL) analysis of a female sexual ornament, the comb of the fowl, Gallus gallus, using a large-scale intercross between red junglefowl and a domestic line, selected for egg production. First, we demonstrate that female somatic investment in comb reflects female reproductive investment. Despite a trade-off between reproductive and skeletal investment mediated by the mobilization of skeletal minerals for egg production, females with proportionally large combs also had relatively high skeletal investment. Second, we identify a major QTL for bisexual expression of comb mass and several QTL specific to female comb mass. Importantly, QTL for comb mass were nonrandomly clustered with QTL for female reproductive and skeletal investment on chromosomes one and three. Together, these results shed light onto the physiological and genetic architecture of a female ornament.

Bivalves of the families Mytilidae, Unionidae, and Veneridae have an unusual mode of mitochondrial DNA (mtDNA) transmission called doubly uniparental inheritance (DUI). A characteristic feature of DUI is the presence of two gender-associated mtDNA genomes that are transmitted through males (M-type mtDNA) and females (F-type mtDNA), respectively. Female mussels are predominantly homoplasmic with only the F-type expressed in both somatic and gonadal tissue; males are heteroplasmic with the M-type expressed in the gonad and F-type in somatic tissue for the most part. An unusual evolutionary feature of this system is that an mt genome with F-coding sequences occasionally invades the male route of inheritance (i.e., a “role reversal” event), and is thereafter transmitted as a new M-type. Phylogenetic studies have demonstrated that the new or “recently masculinized” M-types may eventually replace the older or “standard” M-types over time. To investigate whether this replacement process could be due to an advantage in sperm swimming behavior, we measured differences in motility parameters and found that sperm with the recently masculinized M-type had significantly faster curvilinear velocity and average path velocity when compared to sperm with standard M-type. This increase in sperm swimming speed could explain the multiple evolutionary replacements of standard M-types by masculinized M-types that have been hypothesized for the mytilid lineage. However, our observations do not support the hypothesis that DUI originated because it permits the evolution of mitochondrial adaptations specific to sperm performance, otherwise, the evolutionarily older, standard M genome should perform better.

The genetic incompatibility avoidance hypothesis as an explanation for the polyandrous mating strategies (mating with more than one male) of females of many species has received significant attention in recent years. It has received support from both empirical studies and a meta-analysis, which concludes that polyandrous females enjoy increased reproductive success through improved offspring viability relative to monandrous females. In this study we investigate whether polyandrous female Drosophila simulans improve their fitness relative to monandrous females in the face of severe Wolbachia-associated reproductive incompatibilities. We use the results of this study to develop models that test the predictions that Wolbachia should promote polyandry, and that polyandry itself may constrain the spread of Wolbachia. Uniquely, our models allow biologically relevant rates of incompatibility to coevolve with a polyandry modifier allele, which allows us to evaluate the fate of the modifier and that of Wolbachia. Our empirical results reveal that polyandrous females significantly reduce the reproductive costs of Wolbachia, owing to infected males being poor sperm competitors. The models show that this disadvantage in sperm competition can inhibit or prevent the invasion of Wolbachia. However, despite the increased reproductive success obtained by polyandrous females, the spread of a polyandry modifier allele is constrained by any costs that might be associated with polyandry and the low frequency of incompatible matings when Wolbachia has reached a stable equilibrium. Therefore, although incompatibility avoidance may be a benefit of polyandry, our findings do not support the hypothesis that genetic incompatibilities caused by Wolbachia promote the evolution of polyandry.

In many species of social Hymenoptera, totipotency of workers induces potential conflicts over reproduction. However, actual conflicts remain rare despite the existence of a high reproductive skew. One of the current hypotheses assumes that conflicts are costly and thus selected against. We studied the costs of conflicts in 20 colonies of the queenless ant Diacamma sp. “nilgiri” by testing the effects of conflicts on labor and worker immunocompetence, two parameters closely linked to the indirect fitness of workers. In this species, the dominant female is the only mated worker (gamergate) and monopolizes reproduction. We experimentally induced conflicts by splitting each colony into two groups, a control group containing the gamergate and an orphaned group displaying aggressions until a new dominant worker arises. Immunocompetence was assessed by the clearance of Escherichia coli bacteria that we injected into the ants. Time budget analysis revealed a lower rate of labor and especially brood care in orphaned groups, supporting the existence of a cost of conflicts on labor. Fifteen days after splitting, a lower immunocompetence was also found in orphaned groups, which concerned workers involved and not involved in conflicts. We propose that this immunosuppression induced by conflicts could stem from stress and not directly from aggression.

Demographic parameters including operational sex ratio (OSR) and population density may influence the opportunity for, and strength of sexual selection. Traditionally, male-biased OSRs and high population densities have been thought to increase the opportunity for sexual selection on male sexual traits due to increased male competition for mates. Recent experimental evidence, however, suggests that male-biased OSRs might reduce the opportunity for sexual selection due to increased sexual coercion experienced by females. How OSR, density, and any resultant changes in the opportunity for sexual selection actually affect selection on male sexual traits is unclear. In this study, we independently manipulated OSR and density in the guppy (Poecilia reticulata) without altering the number of males present. We recorded male and female behavior and used DNA microsatellite data to assign paternity to offspring and estimate male reproductive success. We then used linear selection analyses to examine the effects of OSR and density on directional sexual selection on male behavioral and morphological traits. We found that females were pursued more by males in male-biased treatments, despite no change in individual male behavior. There were no differences in sexual behavior experienced by females or performed by males in relation to density. Neither OSR nor density significantly altered the opportunity for sexual selection. Also, Although there was significant multivariate linear selection operating on males, neither OSR nor density altered the pattern of sexual selection on male traits. Our results suggest that differences in either OSR or density (independent of the number of males present) are unlikely to alter directional evolutionary change in male sexual traits.

The adaptive function of female extrapair mating in socially monogamous passerines is currently debated. In the bluethroat (Luscinia s. svecica), a previous study showed that offspring sired by extrapair males had a higher cell-mediated immunity than their within-pair half siblings, suggesting an immunogenetic benefit of extrapair mating in this species. Here, we expanded that dataset with two more years and investigated the association between extrapair paternity and microsatellite multilocus heterozygosity, in addition to cell-mediated immunity. We found that extrapair offspring were more heterozygous than their within-pair half siblings, and corroborated the previous finding of enhanced cellular immunity in extrapair offspring in this four-year dataset. The increased heterozygosity among extrapair offspring appeared to be a result of extrapair mates being less genetically similar than pair mates, and also less genetically similar than expected by random choice. Together with previous findings in this species, showing that the majority of females participate in extrapair copulations, our results indicate a postcopulatory cryptic female choice of genetically dissimilar males. The enhanced cellular immunity and increased heterozygosity were not related to each other, and hence our results indicate two independent genetic benefits of extrapair mating in the bluethroat.

Hermaphroditism allows considerable scope for contributing genes to subsequent generations through various mixtures of selfed and outcrossed offspring. The fitness consequences of different family compositions determine the evolutionarily stable mating strategy and depend on the interplay of genetic features, the nature of mating, and factors that govern offspring development. This theoretical article considers the relative contributions of these influences and their interacting effects on mating-system evolution, given a fixed genetic load within a population. Strong inbreeding depression after offspring gain independence selects for exclusive outcrossing, regardless of the intensity of predispersal inbreeding depression, unless insufficient mating limits offspring production. The extent to which selfing evolves under weak postdispersal inbreeding depression depends on predispersal inbreeding depression and the opportunity for resource limitation of offspring production. Mixed selfing and outcrossing is an evolutionarily stable strategy (ESS) if selfed zygotes survive poorly, but selfed offspring survive well, and maternal individuals produce enough “extra” eggs that deaths of unviable outcrossed embryos do not impact offspring production (reproductive compensation). Mixed mating can also be an ESS, despite weak lifetime inbreeding depression, if self-mating reduces the number of male gametes available for outcrossing (male-gamete discounting). Reproductive compensation and male-gamete discounting act largely independently on mating-system evolution. ESS mating systems always involve either complete fertilization or fertilization of enough eggs to induce resource competition among embryos, so although reproductive assurance is adaptive with insufficient mating, it is never an ESS. Our results illustrate the theoretical importance of different constraints on offspring production (availability of male gametes, egg production, and maternal resources) for both the course and outcome of mating-system evolution, whereas unequal competition between selfed and outcrossed embryos has limited effect. These results also underscore the significance of heterogeneity in the nature and intensity of inbreeding depression during the life cycle for the evolution of hermaphrodite mating systems.

Recent studies have increasingly implicated deep (pre-Pleistocene) events as key in the vertebrate speciation, downplaying the importance of more recent (Pleistocene) climatic shifts. This work, however, has been based almost exclusively on evidence from molecular clock inferences of splitting dates. We present an independent perspective on this question, using ecological niche model reconstructions of Pleistocene Last Glacial Maximum (LGM) potential distributions for the Thrush-like Mourner (Schiffornis turdina) complex in the neotropics. LGM distributional patterns reconstructed from the niche models relate significantly to phylogroups identified in previous molecular systematic analyses. As such, patterns of differentiation and speciation in this complex are consistent with Pleistocene climate and geography, although further testing will be necessary to establish dates of origin firmly and unambiguously.

Early observers of plant evolution in the Arctic have noted a floristic similarity with temperate alpine regions and a predominance of high ploidy levels. The aim of our study was to survey these and other traits in multiple closely related but independently evolved lineages of Artemisia. Our phylogenetic study was based on 133 taxa using 3′-ETS and ITS, and on data on morphology, karyology, distribution, and ecological preferences. We compared Arctic lineages with sister groups and tested whether patterns were significantly different. We found: (1) Artemisia has independently adapted to Arctic habitats 13–18 times; (2) There were no ecological preferences of putative progenitors that might determine the colonization success in the Arctic, although most sister groups were centered in steppe habitats; (3) Plant height was distinctly reduced in Arctic lineages; (4) Arctic lineages contained no more polyploids than their respective sister groups or taxa from other habitats; (5) Enlarged flower heads have evolved repeatedly, probably for better pollinator attraction. This strategy could be a substitute for polyploidy, which is typical in other Arctic taxa. Stronger pollinator attraction should result in better outcrossing and higher heterozygosity in the offspring, which is among the main effects of polyploidy.

Pleiotropy is an aspect of genetic architecture underlying the phenotypic covariance structure. The presence of genetic variation in pleiotropy is necessary for natural selection to shape patterns of covariation between traits. We examined the contribution of differential epistasis to variation in the intertrait relationship and the nature of this variation. Genetic variation in pleiotropy was revealed by mapping quantitative trait loci (QTLs) affecting the allometry of mouse limb and tail length relative to body weight in the mouse-inbred strain LG/J by SM/J intercross. These relationship QTLs (rQTLs) modify relationships between the traits affected by a common pleiotropic locus. We detected 11 rQTLs, mostly affecting allometry of multiple bones. We further identified epistatic interactions responsible for the observed allometric variation. Forty loci that interact epistatically with the detected rQTLs were identified. We demonstrate how these epistatic interactions differentially affect the body size variance and the covariance of traits with body size. We conclude that epistasis, by differentially affecting both the canalization and mean values of the traits of a pleiotropic domain, causes variation in the covariance structure. Variation in pleiotropy maintains evolvability of the genetic architecture, in particular the evolvability of its modular organization.

The genetic basis of sexual dimorphisms is an intriguing problem of evolutionary genetics because dimorphic traits are limited to one sex. Such traits can arise genetically in two ways. First, the alleles that cause dimorphisms could be limited in expression to only one sex at their first appearance. Alternatively, dimorphism alleles could initially be expressed in both sexes, but subsequently be repressed or promoted in only one sex by the evolution of modifier genes or regulatory elements. We investigated these alternatives by looking for the expression of sexually dimorphic traits in female hybrids between bird species whose males show different types of ornaments. If modifier alleles or regulatory elements involved in sex-limited traits are not completely dominant, the modification should break down in female hybrids, which might then show dimorphic traits resembling those seen in males. Of 13 interspecific hybridizations examined, we found not a single instance of the expression of male-limited ornaments in female hybrids. This suggests that male ornaments were sex limited from the outset or that those traits became sex limited through the evolution of dominant modifiers—possibly cis-dominant regulatory elements. Observing hybrid phenotypes is a useful approach to studying the genetics and evolution of dimorphic traits.

Although coevolution is widely accepted as a concept, its importance as a driving factor in biological diversification is still being debated. Because coevolution operates mainly at the population level, reciprocal coadaptations should result in trait covariation among populations of strongly interacting species. A long-tongued fly (Prosoeca ganglbaueri) and its primary floral food plant (Zaluzianskya microsiphon) were studied across both of their geographical ranges. The dimensions of the fly's proboscis and the flower's corolla tube length varied significantly among sites and were strongly correlated with each other. In addition, the match between tube length of flowers and tongue length of flies was found to affect plant fitness. The relationship between flower tube length and fly proboscis length remained significant in models that included various alternative environmental (altitude, longitude, latitude) and allometric (fly body size, flower diameter) predictor variables. We conclude that coevolution is a compelling explanation for the geographical covariation in flower depth and fly proboscis length.

Recent theoretical models and empirical studies of fruit flies, birds, and fish indicate that assortative mating may initiate speciation when physical barriers to gene flow are absent, and before postzygotic barriers evolve. These are important results for marine animals like coral reef fish, where ocean currents can carry planktonic larvae over broad ranges, interconnecting populations and slowing genetic divergence. The Caribbean hamlets (genus Hypoplectrus) are a flock of reef fish morphospecies with highly distinct color pattern that mate like with like, but show little mitochondrial or microsatellite DNA differentiation. Here, we broadly screen genomic diversity using amplified fragment length polymorphisms (AFLP) and survey mating pair formation between two morphospecies in the Florida Keys, the butter hamlet (H. unicolor) and the blue hamlet (H. gemma). No AFLP was species-diagnostic (fixed), and neighbor-joining analyses revealed no clustering of individuals consistent with morphospecies boundaries. Assignment tests, however, placed most individuals within their morphospecies of origin. Field surveys showed that > 98% of mating pairs, including those of rare morphospecies, were of like color pattern. Spawning by a single mixed pair adds to earlier observations suggesting that infrequent hybridization may be a genetically homogenizing force in Hypoplectrus. This study provides a clear example of strong assortative mating in a system with limited genetic differentiation.

The evolution of sex determination remains one of the most fascinating enigmas in biology. Transitions between genotypic sex determination (GSD) and temperature-dependent sex determination (TSD) have occurred multiple times during vertebrate evolution, however, the molecular basis and consequences of these transitions in closely related taxa remain unresolved. Here I address a critical question: Do species with GSD derived from ancestors possessing TSD retain any ancestral thermal sensitivity in the developmental pathways underlying gonadal differentiation? Results from an expression study of a gene involved in early gonadogenesis in GSD (Apalone mutica) and TSD (Chrysemys picta) turtles, support the hypothesis that Wt1 in A. mutica displays such a relic thermal sensitivity. This retention is likely enabled by Sf1, a gene immediately downstream from Wt1 whose expression is independent of temperature in this species. My results constitute the first empirical evidence of a GSD vertebrate exhibiting thermal sensitivity in the expression of a gene regulating gonadogenesis. This novel finding reveals an undocumented source of raw material for future evolutionary change that may exist in other GSD taxa, and one that enhances the evolutionary potential of the gene networks underlying sexual differentiation and contributes to the astonishing ability of sex-determining mechanisms.

The analysis of diversification and character evolution using phylogenetic data attracts increasing interest from biologists. Recent statistical developments have resulted in a variety of tools for the inference of macroevolutionary processes in a phylogenetic context. In a recent paper Maddison (2006 Evolution, 60: 1743–1746) pointed out that uncareful use of some of these tools could lead to misleading conclusions on diversification or character evolution, and thus to difficulties in distinguishing both phenomena. I here present guidelines for the analyses of macroevolutionary data that may help to avoid these problems. The proper use of recently developed statistical methods may help to untangle diversification and character change, and so will allow us to address important evolutionary questions.

Poor public perceptions and understanding of evolution are not unique to the developed and more industrialized nations of the world. International resistance to the science of evolutionary biology appears to be driven by both proponents of intelligent design and perceived incompatibilities between evolution and a diversity of religious faiths. We assessed the success of a first-year evolution course at the University of Cape Town and discovered no statistically significant change in the views of students before the evolution course and thereafter, for questions that challenged religious ideologies about creation, biodiversity, and intelligent design. Given that students only appreciably changed their views when presented with “facts,” we suggest that teaching approaches that focus on providing examples of experimental evolutionary studies, and a strong emphasis on the scientific method of inquiry, are likely to achieve greater success. This study also reiterates the importance of engaging with students' prior conceptions, and makes suggestions for improving an understanding and appreciation of evolutionary biology in countries such as South Africa with an inadequate secondary science education system, and a dire lack of public engagement with issues in science.