Assortative mating and gene flow generate clinal phenological variation in trees

ABSTRACT: BACKGROUND: On-going climate change is shifting the timing of bud burst (TBB) of broad leaf and conifer trees in temperate areas, raising concerns about the abilities of natural populations to respond to these shifts. The level of expected evolutionary change depends on the level and distribution of genetic variation of TBB. While numerous experimental studies have highlighted the role of divergent selection in promoting clinal TBB differentiation, we explored whether the observed patterns of variation could be generated by the joint effects of assortative mating for TBB and gene flow among natural populations. We tested this hypothesis using an in silico approach based on quantitative genetic models. RESULTS: Our simulations showed that genetic clines can develop even without divergent selection. Assortative mating in association with environmental gradients substantially shifted the mean genetic values of populations. Owing to assortative mating, immigrant alleles were screened for proximal or distant populations depending on the strength of the environmental cline. Furthermore, we confirmed that assortative mating increases the additive genetic variance within populations. However, we observed also a rapid decline of the additive genetic variance caused by restricted gene flow between neighboring populations resulting from preferential matings between phenologically-matching phenotypes. CONCLUSIONS: We provided evidence that the patterns of genetic variation of phenological traits observed in forest trees can be generated solely by the effects of assortative mating and gene flow. We anticipate that predicted temperature increases due to climate change will further enhance genetic differentiation across the landscape. These trends are likely to be reinforced or counteracted by natural selection if phenological traits are correlated to fitness.     

Assortative mating,sexual selection,and their consequences for gene flow in Littorina

When divergent populations are connected by gene flow, the establishment of complete reproductive isolation usually requires the joint action of multiple barrier effects. One example where multiple barrier effects are coupled consists of a single trait that is under divergent natural selection and also mediates assortative mating. Such multiple-effect traits can strongly reduce gene flow. However, there are few cases where patterns of assortative mating have been described quantitatively and their impact on gene flow has been determined. Two ecotypes of the coastal marine snail, Littorina saxatilis, occur in North Atlantic rocky-shore habitats dominated by either crab predation or wave action. There is evidence for divergent natural selection acting on size, and size-assortative mating has previously been documented. Here, we analyze the mating pattern in L. saxatilis with respect to size in intensively sampled transects across boundaries between the habitats. We show that the mating pattern is mostly conserved between ecotypes and that it generates both assortment and directional sexual selection for small male size. Using simulations, we show that the mating pattern can contribute to reproductive isolation between ecotypes but the barrier to gene flow is likely strengthened more by sexual selection than by assortment.     

Winter Fidelity and Apparent Survival of Lesser Snow Goose Populations in the Pacific Flyway

Abstract The Beringia region of the Arctic contains 2 colonies of lesser snow geese (Chen caerulescens caerulescens) breeding on Wrangel Island, Russia, and Banks Island, Canada, and wintering in North America. The Wrangel Island population is composed of 2 subpopulations from a sympatric breeding colony but separate wintering areas, whereas the Banks Island population shares a sympatric wintering area in California, USA, with one of the Wrangel Island subpopulations. The Wrangel Island colony represents the last major snow goose population in Russia and has fluctuated considerably since 1970, whereas the Banks Island population has more than doubled. The reasons for these changes are unclear, but hypotheses include independent population demographics (survival and recruitment) and immigration and emigration among breeding or wintering populations. These demographic and movement patterns have important ecological and management implications for understanding goose population structure, harvest of admixed populations, and gene flow among populations with separate breeding or wintering areas. From 1993 to 1996, we neckbanded molting birds at their breeding colonies and resighted birds on the wintering grounds. We used multistate mark-recapture models to evaluate apparent survival rates, resighting rates, winter fidelity, and potential exchange among these populations. We also compared the utility of face stain in Wrangel Island breeding geese as a predictor of their wintering area. Our results showed similar apparent survival rates between subpopulations of Wrangel Island snow geese and lower apparent survival, but higher emigration, for the Banks Island birds. Males had lower apparent survival than females, most likely due to differences in neckband loss. Transition between wintering areas was low (<3%), with equal movement between northern and southern wintering areas for Wrangel Island birds and little evidence of exchange between the Banks and northern Wrangel Island populations. Face staining was an unreliable indicator of wintering area. Our findings suggest that northern and southern Wrangel Island subpopulations should be considered a metapopulation in better understanding and managing Pacific Flyway lesser snow geese. Yet the absence of a strong population connection between Banks Island and Wrangel Island geese suggests that these breeding colonies can be managed as separate but overlapping populations. Additionally, winter population fidelity may be more important in lesser snow geese than in other species, and both breeding and wintering areas are important components of population management for sympatric wintering populations.     

Ultrastructural Studies of Microgametogenesis and Macrogametogenesis of Eimeria truncata of the Lesser Snow Goose1

ABSTRACT. Microgamonts and macrogamonts of Eimeria truncata were observed in renal epithelial cells of collecting tubules and ducts and occasionally in macrophages of experimentally infected lesser snow geese (Anser c. caerulescens) beginning 8.5 days post inoculation. Intraparasitophorous vesicles in parasitophorous vacuoles of both types of gamonts appeared to originate in host cell cytoplasm and enter gamonts through micropores by budding of plasmalemma or by pinocytosis. Within the parasite's cytoplasm, the vesicles were broken down in Golgi-associated vacuoles. The surfaces of microgamonts were highly invaginated to facilitate extrusion of numerous microgametes. Formation and maturation of microgametes were similar to those of other eimerian species. Each microgamete had two flagella, a mitochondrion, and a peculiarly shaped electron-dense nucleus that was oval anteriorly in cross section and somewhat dumbbell-shaped posteriorly. A longitudinally arranged inner membrane complex lay between a portion of the mitochondrion and the plasmalemma. About five subpellicular microtubules extended the length of the microgamete body. Macrogametogony differed little from that described in other eimerian species. Type 1 wall-forming bodies (WFB) formed in Golgi complexes early in macrogametogony, and type 2 WFB formed in cisternae of endoplasmic reticulum in intermediate stages of macrogamont development.     

The Lesser Snow Goose on Wrangel Island: Trophic Interactions with Ruminants and Breeding Habitat Selection

The lesser snow goose (Anser caerulescens caerulescens) has been exterminated across a vast area of Eurasia. At present, it is unable to regenerate there, though its population in North America has reached fifteen million. In Eurasia, the only major nesting colony still persists on Wrangel Island, where the geese use the trophic resources together with ruminants. An assessment of the competitive networks and the trophic interactions between the geese and the ruminants was performed. The analysis of the significance of the trophic niche overlap and the competitive advantages of geese in the habitat preferences has proved that the ruminants are stronger competitors for trophic resources than the geese. It has been ascertained that the levels of competition for trophic resources and/or resource shortages are higher across the habitats of most types associated with the goose colony. The level of the competition for trophic resources is lower, and the feed resources are more diverse and abundant in the habitats that are used by the geese after leaving the colony. It can be concluded that the shortage of resources and/or the stronger competitors for trophic resources (the ruminants) cannot prevent restoration of extinct colonies or the formation of new colonies with a recent increase in the size of the goose population on Wrangel Island. The distribution, abundance, and quality of trophic resources and the competition for them with ruminants do not determine the goose choice of a habitat for a nesting colony. The choice depends on the microclimate and, probably, on a range of other factors.     

Assortative mating and the role of phenotypic plasticity in male competition: implications for gene flow among host-associated parasitoid populations

Local adaptation is promoted when habitat or mating preferences reduce gene flow between populations. However, gene flow is not only a function of dispersal but also of the success of migrants in their new habitat. In this study I investigated mating preference in conjunction with phenotypic plasticity using Aphidius parasitoids adapted to different host species. Males actively attempted to assortatively mate, but actual mating outcomes were strongly influenced by the relative size of the adult males. Results are discussed in the context of assortative mating in combination with the success of migrant males in mitigating gene flow between host-associated parasitoid populations.     

Assortative mating and the genetic correlation

Summary The effect of assortative mating on the genetic correlation between traits X and Y is considered. Assortation on trait X changes the magnitude of the genetic correlation but not its sign. There are two situations depending on the signs of the correlation between mates () and of the random mating genetic correlation (): 1) if sign () = sign (), then >, where is the genetic correlation at equilibrium after continued assortation, and 2) if sign () = sign (), then < . However, negative assortative mating is virtually powerless to alter the magnitude of the genetic correlation. The consequences of a mixed assortation model, e.g., high milk production females mated to fast growing males and lesser productive females mated to slower growing sires, were also studied. Mixed positive assortation always increases the genetic correlation, but negative assortation decreases it. The implications of assortative mating on correlated responses to selection and on the equilibrium covariances between relatives for pairs of traits are discussed.     

Assortative mating by size: A meta-analysis of mating patterns in water striders

Summary Assortative mating by size is a common mating pattern that can be generated by several different behavioural mechanisms, with different evolutionary implications. Assortative mating is typically associated with sexual selection and has been regarded as an attribute of populations, species, mating systems or even higher order taxa. In most animal groups, however, appropriate analyses of assortative mating at these different levels are lacking and the causes and forms of assortative mating are poorly understood. Here, we analyse 45 different population level estimates of assortative mating and non-random mating by size in seven confamiliar species of water striders that share a common mating system. A hierarchical comparative analysis shows that virtually all the variance within the clade occurs among samples within species. We then employ meta-analysis to estimate the overall strength of assortative mating, to determine the form of assortative mating and to further assess potential differences among species as well as the probable causes of assortative mating in this group of insects. We found overall weak but highly significant positive assortative mating. We show that analyses of the degree of heteroscedasticity in plots of male versus female size are critical, since the evolutionary implications of true and apparent assortative mating differ widely and conclude that the positive assortative mating observed in water striders was of the true rather than the apparent form. Further, within samples, mating individuals were significantly larger than non-mating individuals in both males and females. All of these non-random mating patterns were consistent among species and we conclude that weak positive assortative mating by size is a general characteristic of those water strider species that share this mating system. We use our results to illustrate the importance of distinguishing between different forms of assortative mating, to discriminate between various behavioural causes of assortative mating and to assess potential sources of interpopulational variance in estimates of assortative mating. Finally, we discuss the value of using meta-analytic techniques for detecting overall patterns in multiple studies of non-random mating.     

Ultrastructural Studies of the Zygote and Oocyst Wall Formation of Eimeria truncata of the Lesser Snow Goose1

ABSTRACT. Zygote development and oocyst wall formation of Eimeria truncata occurred in epithelial cells in renal tubules and ducts of experimentally infected lesser snow geese (Anser c. caerulescens). Post-fertilization stages were present throughout the kidneys beginning nine days post-inoculation. Initially, a single plasmalemma enclosed the zygote, and type 1 wall-forming bodies (WF1) became labyrinthine and moved toward the surface. There, WF1 degranulated and formed the outer layer of the oocyst wall between the plasmalemma and a newly formed second subpellicular membrane. Several WF2 fused and formed the inner layer, of the oocyst wall between the third and fourth subpellicular membranes. Six subpellicular membranes were observed during wall formation. Other features of oocyst development were similar to those of other eimerian species.     

Assortative mating and the evolution of desirability covariation

Mate choice lies close to differential reproduction, the engine of evolution. Patterns of mate choice consequently have power to direct the course of evolution. Here we provide evidence suggesting one pattern of human mate choice—the tendency for mates to be similar in overall desirability—caused the evolution of a structure of correlations that we call the d factor. We use agent-based models to demonstrate that assortative mating causes the evolution of a positive manifold of desirability, d, such that an individual who is desirable as a mate along any one dimension tends to be desirable across all other dimensions. Further, we use a large cross-cultural sample with n = 14,478 from 45 countries around the world to show that this d-factor emerges in human samples, is a cross-cultural universal, and is patterned in a way consistent with an evolutionary history of assortative mating. Our results suggest that assortative mating can explain the evolution of a broad structure of human trait covariation.     

Genetic modelling: an analysis of a colour polymorphism in the Snow Goose (Anser caerulescens)

Previous studies of colour polymorphism in the Snow Goose [ Anser caerulescens ; failed to consider and reject alternative hypotheses to that of a single locus with incomplete dominance. Utilizing data from a long-term study of a wild population, we test the validity of these earlier results by considering two alternatives: (1) a single locus with multiple allelism and (2) a threshold polygenic system. Our analyses corroborate the original model, but emphasize the importance of testing all plausible hypotheses.     

Assortative versus selective mating: is the distinction worthwhile?

The study examines the distinction between assortative and selective mating made by Lewontin, Kirk, and Crow in 1968 and finds it unproductive. Not only has the difference been ignored on many occasions even as it was invoked, but maintaining it obscures several useful properties of both nonrandom mating schemes and some formally equivalent systems such as fertility selection. The elucidation of these similarities could have accelerated the work of population biologists.