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Unit 1: Evolution chapter 22: descent with modification chapter 25.2 & 3 & 4 : history of life on earth chapter 23: evolution of populations chapter 24 and 25.6 : origin of species chapter 26.1 and 26.3: phylogeny Chapter 24: Origin of Species How do we know that certain organisms belong to the same or different species? • microevolution = change in allele frequencies in a population (caused by mutation, natural selection, genetic drift and flow, etc.) • macroevolution = broad pattern of evolution above species level – Results in origin of new groups (e.g., mammals, flowering plants) through a series of speciation events • speciation = – process by which one species splits into two or more species – bridge between micro- and macroevolution What is a species? Biological species concept: A species is a group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring. The focus is on reproductive compatibility. There are other species concepts (morphological, ecological, phylogenetic, etc.) that emphasize other aspects of species identity. (pg. 504) How are new species formed or maintained? • With gene flow… – populations interbreed, share alleles – maintains a single species • Without gene flow... – populations no longer exchange alleles – genetic changes occur independently (genetic isolation) • Speciation requires reproductive isolation (according to biological species concept). – barriers to interbreeding (prezygotic and postzygotic) – blocks gene flow – maintains separate species Reproductive barriers (Fig. 24.3) prezygotic barriers • habitat isolation Prevent fertilization by: - Impeding the act of mating - Preventing attempted mating from being completed successfully - Specifically hindering the occurrence of fertilization if mating is completed – same geographic region, different habitat water-bound terrestrial • temporal isolation – breed at different times - Impede the act of mating - Prevent attempted mating from being completed successfully - Specifically hinder the occurrence of fertilization if mating is completed • behavioral isolation – different courtship rituals • mechanical isolation – mating is attempted but not completed - Impede the act of mating - Prevent attempted mating from being completed successfully - Specifically hinder the occurrence of fertilization it mating is completed • gametic isolation – sperm cannot fertilize egg - Impede the act of mating - Prevent attempted mating from being completed successfully - Specifically hinder the occurrence of fertilization it mating is completed postzygotic barriers Mating attempt completed successfully and fertilization achieved; prevent hybrid from surviving/thriving and/or reproducing • reduced hybrid viability – offspring of members of different species have lower survival • reduced hybrid fertility – offspring of members of different species are sterile male donkey female horse sterile mule • hybrid breakdown – first generation between two species is viable and fertile – second generation is not viable and fertile “Biological species concept” has limitations… • fossils • asexual organisms • Some species are distinct despite (very limited) gene flow. More than 20 species definitions exist, including… Morphological species concept • distinguishes species by body structure • works for asexual and sexual species • actually used more in practice • very subjective Ecological species concept • distinguishes species by organismal niche in environment • works for asexual and sexual species Phylogenetic species concept • distinguishes species by the smallest group of individuals which share a common ancestor • a species represents one branch of the tree • determining degree of difference can be complicated Speciation can happen with geographic separation… • allopatric speciation – a population is divided into geographically isolated populations – gene flow is limited or eliminated What is the process? • Start with one species, one population • Geographic separation occurs. • Gene flow stops. • One species, two populations • Genetic change occurs independently in each population. – different mutations – natural selection – genetic drift • If the genetic changes lead to the establishment of reproductive isolation mechanisms in these populations, a speciation event can occur. • Note: Geographic isolation alone is not enough for speciation. Genetic changes in the isolated populations must give rise to reproductive isolation in order for speciation to occur. Example of allopatric speciation following the rise of the Panamanian land bridge Speciation can happen without geographic separation… • sympatric speciation – occurs in populations that live in the same area – less common – gene flow is reduced by: • polyploidy – cell division accident results in additional set of chromosomes – more common in plants – oats, cotton, potatoes, tobacco and wheat – can be created in labs • habitat differentiation – a subpopulation is able to exploit a new habitat or resource • sexual selection Fig. 24.14: What happens if species with incomplete reproductive barriers come into contact with one another? How quickly does speciation occur? A survey of data from 84 groups of plants and animals gives us some insight into the total time between speciation events: – minimum: 4,000 yrs (cichlids) – maximum: 40 million yrs (some beetles) – average: 6.5 million yrs and rarely less than 500,000 years This has implications for mass extinction recovery times… Evolution is not goal oriented and cannot create perfect organisms. • Natural selection cannot create “perfect” organisms! – Adaptive evolution ≠ perfectly “engineered” match between environment and organism – Adaptive evolution = good (not perfect) fit; dynamic process! • Evolution “tinkers” via gradual modifications of existing structures and variations, and given enough time, these gradual modifications can produce significant changes (i.e., new species)! • Natural selection and mutation are not predictive. – can only improve a structure in current environment – no anticipation of future advantageous adaptations – no “on demand” system of generating novel, useful alleles (must work with what’s available; limited by historical constraints) • Adaptations are often compromises. • Chance, natural selection, and the environment interact. Complex structures can arise (via evolution) from simpler forms. Limpet (simple mollusc) mollusc Pleurotomaria Nautilus marine snail squid