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Patterns of evolution – Chapter 3 Branching or not • Cladogenesis – lineages branch into two or more lines • Anagenesis – evolutionary change in a lineage without branching Evolutionary history and classification • A taxon (named group of organisms) may be: • Monophyletic (e.g. Aves) • Polyphyletic • Paraphyletic (e.g. Pongidae) Pongidae, a paraphyletic group Patterns of evolution • From phylogenetic studies, common patterns of evolution have been identified 1. Pre-existing features • Features of organisms to not arise de novo (from nothing) • Related organisms have homologous characters, but character state may not be homologous • Homology may be inferred by common position and structure, as well as embryology 2. Homoplasy is common • Types of homoplasy • Convergent evolution • Parallel evolution • Evolutionary reversals • Homoplasy indicates adaptiveness of traits Parallel evolution Convergent evolution Evolutionary reversals 3. Rates of character evolution differ • Some characters evolve very quickly, e.g. body size in mammals • Conservative characters change very little • Mosaic evolution is when different characters evolve at different rates within a lineage 4. Evolution is often gradual • Evolution proceeds by small successive changes (gradualism) rather than by large leaps (saltations) • Not all evolution may be gradual (will talk about later) 6. Change is form is often correlated with change in function 7. Similarity between species changes through ontogeny • Von Baer’s Law – Species are often more similar as embryos than as adults 8. Development underlies some common patterns of morphological evolution • A. Individualization • B. Heterochrony • C. Allometry • D. Heterotopy • E. Simplification of morphology within clades A. Individualization • Bodies of many organisms consist of modules that may become distinct individualization B. Heterochrony • Heterochrony is an evolutionary change in the timing or rate of developmental events • Change in the relative timing of development of somatic features versus reproductive features changes features of organisms • Paedomorphosis • Peramorphosis Paedomorphosis Peramorphosis • Peramorphosis is mostly limited to frontal lobe development and caused: • An increase in the area occupied by the frontal lobe (lobe development is extended), which in turn causes • an increase in the area occupied by the stomach, which • allowed descendants to exploit a different ecological niche (in this case an increased sedimentation regime). C. Allometry • Allometry is the differential rate of growth of different parts of an organism during its development Allometry • Allometry can be described by: • Log y = log b a log x, where a is the allometric coefficient or relative growth rate • If a = 1, then growth is isometric Allometry • Suppose that x is body size and that y is the size of some feature that begin to develop at age α and stops growing at age β Allometry Allometry • If development is extended by Δβ, then paramorphosis results Allometry • If growth is shortened by Δβ (progenesis), then paedomorphosis results Allometry • If the growth rate of y is reduced relative to x (neoteny), paedomorphosis results D. Heterotopy • Heterotopy is an evolutionary change in the position within an organism at which a phenotypic character is expressed Sesamoids E. Simplification of morphology 9. Evolutionary trends • Phylogenetic analysis can also document evolutionary trends. i.e., a succession of changes of a character in the same direction, either within a single lineage or in may lineages independently • For example in plants, we see trends from: • Low to high chromosome number • Animal to wind pollination • Radial to bilateral symmetry • Woody to herbaceous 10. Adaptive radiation is common • Adaptive radiation is the divergent evolution of a lineage within a relatively short time • Mammal and angiosperm diversification during Mesozoic and Cenozoic • Cichlid fishes in rift lakes of Africa • Darwin’s Finches on the Galapagos Islands • Honeycreepers in Hawaii What causes adaptive radiations? • 1. Opportunity • Colonization of isolated habitats • Cuts off gene flow • Many new niches available • Lack of competition • Mass extinction • Climate Change • 2. Evolutionary innovation Colonization of isolated habitats Mass extinctions Evolution of land plants includes major innovations Climate change • Why did Conifers replace Ferns and fern allies? • • Carboniferous forest (ca. 330 mya) Lycopsids, Ferns, Horsetails still: • Required water for fertilization • Have free-living gametophyte generation (no dessication –resistant propagule) Why did gymnosperms replace lycophytes, horsetails, & ferns? • • Seeds evolved by end of Devonian, yet non-seed tracheophytes continued to dominate into Permian What changed to favor Gymnosperms? Rise of the gymnosperms • Formation of Pangea dramatically changed continental climate • Single landmass, very dry • Cold, dry periods • Favorable conditions for gymnosperms • Ferns have flagellated sperm & free-living gametophytes • Gymnosperms & Cycads: wind-dispersed pollen (and first animaldispersed pollen), retain seed