Speciation and Macroevolution • Speciation • Divergent Evolution • Convergent Evolution Evolutionary change of a species over time Creation of new groupspeciation diversity of living things. Speciation & Macroevolution • Microevolution = evolution on small scale • changes in allele frequency in a population over time • Speciation = formation of new species • Bridges micro and macro-evolution • Macroevolution = large scale evolution • broad patterns of evolutionary change; species formation and larger What is a species? • biological species concept: • A group of individuals (i.e., multiple pops.) that have the potential to interbreed in nature and produce viable, fertile offspring. • Based on reproductive compatibility and isolation • Weakness of this def: • Other species definitions: • • • • Morphological species concept: based on structure Ecological species concept: based on niche (i.e., role in ecosystem) Phylogenetic species concept: based on common ancestor (evolutionary relationship) More….. Reproductive Isolation • Reproductive isolation : inability to mate and produce viable, fertile offspring • Classification/Types of Isolation: • Prezygotic Barriers • Pre-mating attempt • Post-mating attempt • Post-zygotic Barriers Isolated population diverges. Gene flow Population Barrier to gene flow Figure 24.13-2 Mating doesn’t occur Gametes don’t unite Offspring doesn’t survive or can’t perpetuate species • Prezygotic barriers block fertilization from occurring by • Impeding different species from attempting to mate • Preventing the successful completion of mating • Hindering fertilization if mating is successful • Postzygotic barriers prevent the hybrid zygote from developing into a viable, fertile adult • Reduced hybrid viability • Reduced hybrid fertility • Hybrid breakdown Two forms of Speciation: • Allopatric: speciation with geographic separation • Non-overlapping populations • “Barrier” is relative to dispersal ability of organism • Even if contact is restored, interbreeding is prevented • Sympatric: speciation within the same geographic location/area • Overlapping Allopatric Speciation: A tale of two fuzzy animals Experiment Initial population of fruit flies (Drosophila pseudoobscura) Some flies raised on starch medium Some flies raised on maltose medium Mating experiments after 40 generations Results Female 9 8 20 Number of matings in experimental group Starch population 2 Starch population 1 Starch 22 Starch population 1 18 15 Starch population 2 Maltose Male Starch Maltose Female Male • Reproductive barriers can develop when laboratory populations are experimentally isolated and subjected to different environmental conditions 12 15 Number of matings in control group Figure 24.7 (a) Under high predation (b) Under low predation Females from each pop show strong preference for their own body shape in mate selection Figure 24.6 Sympatric Speciation • Mechanisms for sympatric speciation • Habitat differentiation • Sexual selection • Polyploidy A “tale” of two squirrels Sexual Selection • Preference for specific traits (e.g., morphological or behavioral) by subset of females can lead to reproductive isolation. • E.g. Sexual selection for mates of different colors has likely contributed to speciation in cichlid fish in Lake Victoria I love a man with a red nose Orange noses are dreamy Figure 24.11 Experiment Normal light P. pundamilia P. nyererei Monochromatic orange light Polyploidy • Polyploidy is the presence of extra sets of chromosomes due to accidents during cell division • Polyploidy can produce new biological species in sympatry within a single generation • Autopolyploid: from a single species • Allopolyploid: from hybrids of two species Cell division error • An autopolyploid is an individual with more than two chromosome sets, derived from a single species 2n = 6 Tetraploid cell 4n Meiosis 2n 2n Gametes produced by tetraploids New species (4n) Autopolyploid speciation Figure 24.UN02 • An allopolyploid is a species with multiple sets of chromosomes derived from different species Species A 2n = 6 Normal gamete n=3 Species B 2n = 4 Normal gamete n=2 Sterile hybrid with 5 chromosomes Mitotic or meiotic error doubles the chromosome number. New species: viable, fertile hybrid (allopolyploid) 2n = 10 Figure 24.9-3 Figure 24.10 T. dubius (12) Hybrid species: T. miscellus (24) T. pratensis (12) Hybrid species: T. mirus (24) T. porrifolius (12) Rate of speciation • It can happen quickly (punctuated equilibrium model) • It can happen slowly (gradualism) • The interval between speciation events can range from 4,000 years (some cichlids) to 40 million years (some beetles), with an average of 6.5 million years (a) Punctuated model Time (b) Gradual model Figure 24.16 Studying the Genetics of Speciation • How many genes change when a new species forms? • As few as a single gene or in other cases many genes • For example, in Japanese Euhadra snails, the direction of shell spiral affects mating and is controlled by a single gene • In monkey flowers (Mimulus), two loci affect flower color, which influences pollinator preference • Pollination that is dominated by either hummingbirds or bees can lead to reproductive isolation of the flowers • In other organisms, speciation can be influenced by larger numbers of genes and gene interactions 68x more hummmingbrid visits than wild-type M. lewisii Preferred by bumblebees (a) Mimulus lewisii (b) M. lewisii with M. cardinalis allele 74x more bumblebee visits than wild-type M. cardinalis Preferred by hummingbirds (c) Mimulus cardinalis (d) M. cardinalis with M. lewisii allele Figure 24.19 Convergent Evolution • When similar selective forces cause similarities (similar traits) in unrelated species • E.g., body shape of aquatic organism (shark and dolfins) • Similar structure of desert plants cacti & euphorbs