Speciation • “It is really laughable to see what different ideas are prominent in various naturalists’ minds, when they speak of ‘species’. It all comes, I believe, from trying to define the indefinable”. • Charles Darwin What is a species? Using the BSC • "a species is an array of populations which are actually or potentially interbreeding, and which are reproductively isolated from other such arrays under natural conditions." (Ernst Mayr) • Reproductive isolation is the failure of populations to interbreed or to form viable or fertile hybrids • Speciation = generation of a reproductive isolating mechanism Fig. 24-4 Prezygotic barriers Habitat Isolation Temporal Isolation Individuals of different species (a) Postzygotic barriers Behavioral Isolation Mechanical Isolation Gametic Isolation Mating attempt (c) (d) (e) (f) Reduced Hybrid Viability Reduced Hybrid Fertility Hybrid Breakdown Viable, fertile offspring Fertilization (g) (h) (i) (j) (b) (k) (l) Fig. 24-4a Prezygotic barriers Habitat Isolation Temporal Isolation Individuals of different species (a) Mating attempt (c) (d) (b) Mechanical Isolation Behavioral Isolation (e) (f) Fig. 24-4i Prezygotic barriers Gametic Isolation Postzygotic barriers Reduced Hybrid Viability Reduced Hybrid Fertility Hybrid Breakdown Viable, fertile offspring Fertilization (g) (h) (i) (j) (k) (l) Speciation is often gradual, so we expect to see some “gray areas” (are they different species or not?) Speciation is often gradual, so we expect to see some “gray areas” (are they different species or not?) Canis lycaon Canis latrans Canis lupus Speciation is often gradual, so we expect to see some “gray areas” (are they different species or not?) Canis latrans Canis lycaon Canis Soup Canis lupus The BSC can only apply to a small portion of life The PSC • the smallest set of organisms that share a common ancestor and can be distinguished from other such sets. • Or • the smallest diagnosable cluster of individual organisms within which there is a parental pattern of ancestry and descent (Cracraft 1983) • Speciation = cladogenesis Eastern and Western Northern Flickers • Morphological differences evolved in allopatry • Differences don’t function as premating RIM’s (they hybridize in the Great Plains) • Don’t seem to be any postmating RIM’s (hybrids fit and reproduce) Moore et al., 1991, Mol. Biol. Evol. 8(3):327-344 Types of speciation • Anagenesis—draw it Types of speciation • Anagenesis—draw it • Cladogenesis—draw it Types of speciation • Anagenesis—draw it • Cladogenesis—draw it • Reticulate speciation—draw it Bold = species thought to have formed as hybrids Red arrows = mtDNA source Koblmüller et al. BMC Evolutionary Biology 2007 7:7 doi:10.1186/1471-2148-7-7 3 hybrid species of sunflower that thrive where the parental species cannot