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Using Multivariate Statistics

Statistical method for testing the neutral mutation hypothesis by DNA polymorphism.

So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

The repeatability of behaviour: a meta-analysis.

Developmental Plasticity and Evolution. In: Mary Jane West-Eberhard, Editor, Oxford University Press (2003) Pp. xi+794. Price $50.00 paper.

Nest selection has important consequences for fitness in freshwater turtles, as nest predation is extremely high in most populations. However, nest predation has rarely been considered a major factor affecting nest site selection. Thermal qualities of a nest are often suggested as a factor influencing maternal choice of nest site because incubation temperature affects offspring performance and growth, as well as the sex ratio of the clutch in species with temperature dependent sex determination (TSD). Yet experimental manipulations of the nesting environment (habitat and threat) to test factors influencing maternal nest site selection in turtles are uncommon. In this paper, we show that nest site selection in an Australian turtle (Emydura macquarii) with genetically determined sex determination (GSD) exists at two spatial scales, with predation as the major factor affecting the location of a nest. Females prefer to nest in areas where nest predation is minimal but when the risk of direct predation is increased, they trade-off minimising nest predation with survival by locating their nests closer to shore. However, experimentally reducing cover demonstrated that females prefer to nest in open areas, which are more common closer to shore. Hence females are forgoing preferred microhabitat to locate their nests away from shore to minimise nest predation. Despite a preference for open nesting areas, females exhibit similar repeatability of microenvironment selection to a North American species with TSD. Repeatability of nesting overstory vegetation in the North American turtle, Chrysemys picta, suggests that females may exert some control over the thermal qualities of the nests; a key assumption in theoretical models of the microevolution and adaptive significance of TSD in reptiles. Incubation temperature is one factor that may affect microhabitat preferences in both turtles but other factors (e.g. predation and soil moisture) affected by microhabitat may be critical in the evolution of repeatability of overstory cover.

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The significance of predation in nest site selection of turtles: an experimental consideration of macro- and microhabitat preferences

Squamate reptiles are uniquely suited to study of evolution of reproductive mode and pattern of embryonic nutrition. Viviparous species have evolved from oviparous ancestors on numerous occasions, patterns of nutritional provision to embryos range widely from lecithotrophy, at one end of a continuum, to placentotrophy at the other, and structure and function of the maternal–embryonic relationship is highly constrained resulting in parallel evolutionary trajectories among taxa. Embryos of oviparous species primarily receive nourishment from yolk, but also mobilize a significant quantity of calcium from the eggshell. Most viviparous species also are predominantly lecithotrophic, yet all viviparous species are placentotrophic to some degree. Similarities in embryonic development and nutritional pattern between oviparous species and most viviparous species suggest that the pattern of nutrition of oviparous squamates is an exaptation for the evolution of viviparity and that placentotrophy and viviparity evolve concomitantly. The few species of squamates that rely substantially on placentotrophy have structural modifications of the interface between the embryo and mother that are interpreted as adaptations to enhance nutritional exchange. Recent studies have extended understanding of the diversity of embryonic nutrition and placental structure and have resulted in hypotheses for transitions in the evolution of placentotrophy, yet data are available for few species. Indirect tests of these hypotheses, by comparison of structural–functional relationships among clades in which viviparity has evolved, awaits further study of the reproductive biology of squamates.

Evolution of placentation among squamate reptiles: recent research and future directions.

No one suspected that temperature-dependent sex determination (TSD)1,2,3, whereby the sex of embryos depends on the temperature at which they develop, might occur in viviparous (live-bearing) reptiles, because thermoregulation in the mother results in relatively stable, raised gestation temperatures. But here we show that developing embryos of the actively thermoregulating viviparous skink Eulamprus tympanum are subject to TSD, offering the mother the chance to select the sex of her offspring and a mechanism to help to balance sex ratios in wild populations.

Sex determination. Viviparous lizard selects sex of embryos.

The aim of this review is to collate data relevant to understanding the evolution of viviparity in general, and complex placentae in particular. The wide range of reproductive modes exhibited by lizards provides a solid model system for investigating the evolution of viviparity. Within the lizards are oviparous species, viviparous species that have a very simple placenta and little nutrient uptake from the mother during pregnancy (lecithotrophic viviparity), through a range of species that have intermediate placental complexities and placental nutrient provision, to species that lay microlecithal eggs and most nutrients are provided across the placenta during development (obligate placentotrophy). In its commonest form, lecithotrophic viviparity, some uptake of water, inorganic ions and oxygen occurs from the mother to the embryo during pregnancy. In contrast, the evolution of complex placentae is rare, but has evolved at least five times. Where there is still predominantly a reliance on egg yolk, the omphaloplacenta seems to be paramount in the provision of nutrition to the embryo via histotrophy, whereas the chorioallantoic placenta is more likely involved in gas exchange. Reliance on provision of substantial organic nutrient is correlated with the regional specialisation of the chorioallantoic placenta to form a placentome for nutrient uptake, particularly lipids, and the further development of the gas exchange capabilities of the other parts of the chorioallantois.

A review of the evolution of viviparity in lizards: structure, function and physiology of the placenta

We have reviewed published and new quantitative data on the net uptake of nutrients by embryos of oviparous and viviparous lizards that vary in chorioallantoic placental complexity to better understand the evolution of complex placentae. We assessed net nutrient uptake during embryonic development by measuring the total dry mass, or the mass of separate nutrients, in the egg at about the time of ovulation and in the neonate. There is no significant difference in the fresh egg to neonate dry mass ratio of oviparous and viviparous species that have simple placentae, indicating that there is little, if any, net uptake of nutrients by viviparous species with simple chorioallantoic placentae. In contrast, there is significant uptake of dry matter and individual nutrients across the placenta of species with complex chorioallantoic placentae. Species of the genus Niveoscincus have a range of placentae and nutrient uptakes, even among populations of one species, suggesting that further studies among populations of single species are required. Data are available for relatively few clades, and all the data for the three most complex chorioallantoic placental types are derived from a single genus. Thus, further research on new genera of lizards is required to overcome the potentially confounding effects of phylogeny in our analyses.

Michael B. Thompson

Papers

The significance of predation in nest site selection of turtles: an experimental consideration of macro- and microhabitat preferences

Evolution of placentation among squamate reptiles: recent research and future directions.

Sex determination. Viviparous lizard selects sex of embryos.

A review of the evolution of viviparity in lizards: structure, function and physiology of the placenta

Comparison of nutrient transport across the placenta of lizards differing in placental complexity.