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Origin of species Cladogenesis  One ancestral species becomes divided into two descendant species  If species are defined by the existence of reproductive isolation, then  the process of speciation is identical to the evolution of reproductive isolating mechanisms Reproductive isolation  Populations can become isolated  Geographically in this example Evolution of reproductive isolation  The formation of species is a continuous process  Two populations may only be partially reproductively isolated  If isolating mechanisms have not evolved, then two populations will interbreed freely  If populations are reproductively isolated, no genetic exchange will occur, two populations will be different species Reproductive Isolation  During the time when reproductive isolation is incomplete (the intermediate state):  Hybrids are partly sterile  Hybrids are not as well adapted to the habitat  Selection would favor any alleles in the parental populations that prevent hybridization  Reinforcement - incomplete isolating mechanisms are reinforced by natural selection until they are completely effective Allopatric species  Natural selection led to reproductive isolation where ranges overlapped Gene flow may counter speciation  Reinforcement is not inevitable  Incompletely isolated populations have gene flow  Hybrids may be inferior but serve as a conduit of genetic exchange  Two populations will lose their genetic distinctiveness  A race between complete reproductive isolation evolution and gene flow Genetic drift  Random changes may cause reproductive isolation  Genetic drift in small populations  Founder effects  Population bottlenecks Natural selection  Adaptation can lead to speciation  Natural selection produces a variety of differences in physiological and sensory traits  Promotes ecological and behavioral isolation Anolis dewlap Is geographic isolation required for speciation to occur?  i.e., allopatric speciation Sympatric speciation  Occurs without geographic speciation Polyploidy  Individuals that have more than two sets of chromosomes  e.g., 3n, 4n, etc.  Offspring with altered chromosome number cannot breed with parent population  Common mechanism of speciation in flowering plants  Occurs in insects, fish, and salamanders but is rare Polyploidy  Can occur when all chromosomes arise from a single species (autopolyploidy) or when chromosomes arise from hybridization between two species (allopolyploidy) Autopolyploidy Allopolyploidy Disruptive selection  Sympatric speciation may occur over the course of multiple generations through disruptive selection  Two phenotypes would have to evolve reproductive isolating mechanisms Sympatric speciation in African Cichlid fish  Studied in two lakes in Cameroon  Species in each lake are most likely descended from single ancestor  No barriers within either lake, but 11 species found in one, 9 species in the other  Some ecological isolation by feeding preference Adaptive radiations  Closely related species that have recently evolved from a common ancestor by adapting to different parts of the environment  Occurs in an environment with few other species and many resources  Newly formed islands such as Hawaiian and Galápagos Islands  Catastrophic event leading to extinction of other species Rapid speciation often follows the evolution of a key innovation  A key innovation allows the species possessing it to use resources or other aspects of the environment that were previously inaccessible  Evolution of lungs in fish  Wings in birds and insects  Seeds in plants Adaptive radiation with allopatric speciation Adaptive radiation with sympatric speciation Character displacement  Natural selection in each species favors those individuals that use resources not used by the other species Hawaiian Drosophila  > 1000 species of Drosophila on Hawaiian Islands  Diversity of morphological and behavioral traits  Empty habitats resulted in fruit flies that are:  Predators  Herbivores  Detritivores  Nectar eaters  Parasites Darwin’s finches Lake Victoria cichlids  Was home to over 300 species of cichlid until recently  Recent radiation: sequencing of cytochrome b gene -- 2000,000 years ago  Colonized from the Nile Cichlid extinction  Abrupt extinction in the last several decades  1950’s Nile perch introduced into lake  1990’s 70% cichlids extinct New Zealand alpine buttercups  Speciation in glacial habitats  Periodic isolation  14 species occupy 5 distinct habitats The pace of evolution  Gradualism: the accumulation of small changes  Punctuated equilibrium: long periods of stasis followed by rapid change  Proposed by Niles Eldredge and Stephen Gould in 1972  Stabilizing and oscillating selection is responsible for stasis Gradualism or punctuated equilbrium – which is correct?  Evolution may include both types of change  Rapid change and speciation are not linked  Speciation can occur without substantial phenotypic change  Phenotypic change can occur within species in the absence of speciation Speciation and extinction  Speciation, through time, has surpassed extinction  Five mass extinctions have occurred  Most severe at the end of the Permian period—96% of all species may have perished  End of the Cretaceous – K-T extinction  Dinosaurs went extinct Consequence of extinction  previously dominant groups may perish, changing the course of evolution  Dinosaurs went extinct, mammals began their radiation  Rates of speciation after an extinction may take about 10 my  Takes time for:  Ecosystems to recover  Processes of speciation and adaptive diversification to begin  Not all groups of organisms are affected equally during extinctions A sixth extinction is underway  Estimates:  1/4th of all species will become extinct in the near future  Rebound in species diversity may be slower than following previous mass extinction events  A large proportion of the world’s resources will be taken up by human activities The future of evolution  Human influences on the environment affect the evolutionary processes  Changing patterns of natural selection  Global climate change is major challenge for many species  Decreased population sizes will increase the likelihood of genetic drift  Geographic isolation will remove homogenizing effect of gene flow  Chemicals and radiation could increase mutation rate Tigers now exist in geographically isolated populations