Increased Floral Rewards due to Local Adaptation Drives Plant Ecological Speciation via Learned Preferences of Pollinators.

AbstractIn animal-pollinated plants, the growth environment and pollination environment are two important agents of natural selection. However, their simultaneous effects on plant speciation remain underexplored. Here, we report a theoretical finding that if plants' local adaptation to the growth environment increases their floral rewards for pollinators, it can strongly facilitate ecological speciation in plants. We consider two evolving plant traits, vegetative and floral signal traits, in a population genetic model for two plant populations under divergent selection from different growth environments. The vegetative trait determines plants' local adaptation. Locally adapted plants reward pollinators better than maladapted plants. By associative learning, pollinators acquire learned preferences for floral signal traits expressed by better-rewarding plants. If pollinators' learned preferences become divergent between populations, floral signal divergence occurs and plants develop genetic associations between vegetative and floral signal traits, leading to ecological speciation via a two-allele mechanism. Interestingly, speciation is contingent on whether novel floral signal variants arise before or after plant populations become locally adapted to the growth environment. Our results suggest that simultaneous selection from growth and pollination environments might be important for the ecological speciation of animal-pollinated plants.

The propagation of admixture-derived adaptive radiation potential.

Adaptive radiations (ARs) frequently show remarkable repeatability where single lineages undergo multiple independent episodes of AR in distant places and long-separate time points. Genetic variation generated through hybridization between distantly related lineages can promote AR. This mechanism, however, requires rare coincidence in space and time between a hybridization event and opening of ecological opportunity, because hybridization generates large genetic variation only locally and it will persist only for a short period. Hence, hybridization seems unlikely to explain recurrent AR in the same lineage. Contrary to these expectations, our evolutionary computer simulations demonstrate that admixture variation can geographically spread and persist for long periods if the hybrid population becomes separated into isolated sub-lineages. Subsequent secondary hybridization of some of these can reestablish genetic polymorphisms from the ancestral hybridization in places far from the birthplace of the hybrid clade and long after the ancestral hybridization event. Consequently, simulations revealed conditions where exceptional genetic variation, once generated through a rare hybridization event, can facilitate multiple ARs exploiting ecological opportunities available at distant points in time and space.

Pregnancy parturition scars in the preauricular area and the association with the total number of pregnancies and parturitions.

OBJECTIVES: The aim of the present study was to clarify the association between the degree of development of pregnancy parturition scars (PPSs) and the total number of pregnancies and parturitions (TNPPs) on the basis of new identification standards for PPS in the preauricular area. MATERIALS AND METHODS: Preauricular grooves were macroscopically observed on the pelves of 103 early modern males and 295 females (62 early modern females; 233 present-day females). Three categories of PPS in the preauricular area were defined. The association between the degree of development of PPS in the preauricular area and the TNPP was analyzed in 90 present-day females with detailed lifetime data. RESULTS: PPS could not estimate the exact TNPP. However, it was shown that no PPS indicated no TNPP, weak PPS indicated a lower TNPP, and developed PPS indicated a higher TNPP. DISCUSSION: Even though the possibility remains that some PPS indicate no TNPP, the results showed that the percentage of each PPS category indicated fertility in the population, suggesting that the strength of the association between the degree of development of PPS and the TNPP was affected by the classification system, the reliability of lifetime data, and the statistical methods used for analysis.

Hybridization can promote adaptive radiation by means of transgressive segregation.

Understanding the mechanisms of rapid adaptive radiation has been a central problem of evolutionary ecology. Recently, there is a growing recognition that hybridization between different evolutionary lineages can facilitate adaptive radiation by creating novel phenotypes. Yet, theoretical plausibility of this hypothesis remains unclear because, for example, hybridization can negate pre-existing species richness. Here, we theoretically investigate whether and under what conditions hybridization promotes ecological speciation and adaptive radiation using an individual-based model to simulate genome evolution following hybridization between two allopatrically evolved lineages. The model demonstrated that transgressive segregation through hybridization can facilitate adaptive radiation, most powerfully when novel vacant ecological niches are highly dissimilar, phenotypic effect size of mutations is small and there is moderate genetic differentiation between parental lineages. These results provide a theoretical basis for the effect of hybridization facilitating adaptive radiation.

Inaccurate Color Discrimination by Pollinators Promotes Evolution of Discrete Color Polymorphism in Food-Deceptive Flowers.

Many plant species employing a food-deceptive pollination strategy show discrete or continuous floral polymorphism within their populations. Previous studies have suggested that negative frequency-dependent selection (NFDS) caused by the learning behavior of pollinators was responsible for the maintenance of floral polymorphism. However, NFDS alone does not explain why and when discrete or continuous polymorphism evolves. In this study, we use an evolutionary simulation model to propose that inaccurate discrimination of flower colors by pollinators results in evolution of discrete flower color polymorphism. Simulations showed that associative learning based on inaccurate discrimination in pollinators caused disruptive selection of flower colors. The degree of inaccuracy determined the number of discrete flower colors that evolved. Our results suggest that animal behavior based on inaccurate discrimination may be a general cause of disruptive selection that promotes discrete trait polymorphism.

Predation on pollinators promotes coevolutionary divergence in plant-pollinator mutualisms.

Coevolution of plants and pollinators has been suggested as a mechanism driving diversification of plant-pollinator mutualisms. There is increasing recognition that predators or competitors can influence the abundance and behavior of pollinators and indirectly affect the fitness of plants. However, existing theories on plant-pollinator diversification focus exclusively on mutualistic interactions between plants and pollinators. Here we used simulations to evaluate whether predation on pollinators promotes coevolutionary diversification of plant-pollinator mutualisms. We developed an individual-based simulation model in which the blooming season of plants and the active seasons of pollinators and predators can evolve. In simulations without predators, plant-pollinator coevolution caused diversification in blooming/active seasons for both plants and pollinators, but this diversification resulted in polymorphisms, not speciation. The introduction of predators promoted a split of plant and pollinator populations into reproductively isolated subpopulations with corresponding blooming and active seasons or a directional shift of blooming and active seasons, increasing the possibility of plant-pollinator cospeciation. This result suggests that predation on pollinators can promote sympatric and allopatric divergence of plant-pollinator mutualisms. Joint action of antagonistic and mutualistic interactions may be fundamentally important for diversification in coevolutionary interactions.

Transgressive segregation in mating traits drives hybrid speciation.

Hybridization can rapidly generate novel genetic variation, which can promote ecological speciation by creating novel adaptive phenotypes. However, it remains unclear how hybridization, creating novel mating phenotypes (e.g., mating season, genitalia shapes, sexual displays, mate preferences), affects speciation especially when the phenotypes do not confer adaptive advantages. Here, based on individual-based evolutionary simulations, we propose that transgressive segregation of mating traits can drive incipient hybrid speciation. Simulations demonstrated that incipient hybrid speciation occurred most frequently when the hybrid population received moderate continued immigration from parental lineages causing recurrent episodes of hybridization. Recurrent hybridization constantly generated genetic variation, which promoted the rapid stochastic evolution of mating phenotypes in a hybrid population. The stochastic evolution continued until a novel mating phenotype came to dominate the hybrid population, which reproductively isolates the hybrid population from parental lineages. However, too frequent hybridization rather hindered the evolution of reproductive isolation by inflating the variation of mating phenotypes to produce phenotypes allowing mating with parental lineages. Simulations also revealed conditions for the long-term persistence of hybrid species after their incipient emergence. Our results suggest that recurrent transgressive segregation of mating phenotypes can offer a plausible explanation for hybrid speciation and radiations that involved little adaptive ecological divergence.

Evolution of increased phenotypic diversity enhances population performance by reducing sexual harassment in damselflies.

The effect of evolutionary changes in traits and phenotypic/genetic diversity on ecological dynamics has received much theoretical attention; however, the mechanisms and ecological consequences are usually unknown. Female-limited colour polymorphism in damselflies is a counter-adaptation to male mating harassment, and thus, is expected to alter population dynamics through relaxing sexual conflict. Here we show the side effect of the evolution of female morph diversity on population performance (for example, population productivity and sustainability) in damselflies. Our theoretical model incorporating key features of the sexual interaction predicts that the evolution of increased phenotypic diversity will reduce overall fitness costs to females from sexual conflict, which in turn will increase productivity, density and stability of a population. Field data and mesocosm experiments support these model predictions. Our study suggests that increased phenotypic diversity can enhance population performance that can potentially reduce extinction rates and thereby influence macroevolutionary processes.