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The Origin of Species Chapter 24 BCOR 012 Jan. 31 and Feb. 2, 2011 Outline for January 31, Feb. 2, 2011: I. Introduction II. What is a species? The biological species concept Reproductive isolating mechanisms Alternative species concepts III. Modes of speciation Allopatric Sympatric the tempo of change What is a species? The Biological Species Concept The biological species concept defines a species as a population or a series of populations whose members have the potential to interbreed and which are reproductively isolated from other such populations. Reproductive Isolation and Isolating Mechanisms ex. fireflies Biologists distinguish between prezygotic and postzygotic isolating mechanisms. ex:mule Limitations of the BSC: • Not applicable to asexually reproducing organisms • Useless with respect to the fossil record Alternative species concepts: • Ecological species concept - defines a species in terms of its ecological niche • Morphological species concept - emphasis is on unique structural features • Phylogenetic species concept - emphasis is on ancestor-descendent relationships • Pluralistic species concept - acknowledges that, where species concepts are concerned, one size may not fit all! There are an awful lot of living things; how did there get to be so many? Anagenesis is change within a lineage; cladogenesis is the divergence of one lineage into two. Modes of Speciation There are two general modes of speciation: • Allopatric (other homeland)- speciation takes place in populations with geographically separate ranges • Sympatric (same homeland) - speciation takes place in geographically overlapping populations Ammospermophilus harrisi S Ammospermophilus leucurus N Allopatric speciation of antelope squirrels in the Grand Canyon In allopatric speciation, a new species originates while geographically isolated from its ancestor. As the new species evolves by genetic drift and natural selection, reproductive isolation from the ancestral species may evolve as a by-product of genetic change. Sympatric speciation - a mode in which a new species arises in the geographic midst of its progenitor species. Copper-tolerant Mimulus guttatus In plants, sympatric species may arise by polyploidy a condition that results in extra sets of chromosomes in the derivative species. An example from the European holly ferns …. European holly ferns (Polystichum) P. setiferum P. lonchitis P. aculeatum The origin of European Polystichum aculeatum P. aculeatum (4X) P. setiferum (2X) primary diploid hybrid P. lonchitis (2X) The origin of European Polystichum aculeatum P. aculeatum P. setiferum P. setiferum n=41 primary diploid hybrid P. lonchitis Sympatric speciation in animals East African Cichlids (sick-lids) Pundamilia pundamilia Pundamilia nyererei a) Normal light b) Monochromatic orange light Non-random mating, in which females select mates having a preferred appearance, is the main reproductive barrier keeping these two species separate in nature. Thi is an example of sympatric speciation in response to sexual selection. Sympatric speciation requires the emergence of some sort of reproductive barrier that isolates the gene pool of a population subset without geographic separation from the parent population. Hybrid Zones The Tempo of Evolution Darwin’s original idea was that species diverge gradually. More recently, the idea that there are rapid speciation events interrupting periods of no change has become popular. Punctuated equilibria vs. gradual divergence http://evolution.berkeley.edu/evosite/evo101/VIIA1bPunctuated.shtml Punctuated equilibria vs. gradual divergence http://evolution.berkeley.edu/evosite/evo101/VIIA1bPunctuated.shtml Possibly include spur story Pollinator shifts drive increasingly long nectar spurs in columbine flowers. Hodges, 2007 ---- Nature 447: 706-710. Whittall and Outline for January 31, Feb. 2, 2011: I. Introduction II. What is a species? The biological species concept Reproductive isolating mechanisms Alternative species concepts III. Modes of speciation Allopatric Sympatric the tempo of change