Species and Speciation Organisms • Organisms are classified into species • Number of species on earth: 5-100 million (only ~1.4 million species described ) • 70% of the world's species are in only 12 countries: Australia, Brazil, China, Colombia, Ecuador, India, Indonesia, Madagascar, Mexico, Peru, and Zaire. Species A Species B Non-Evolutionary Approaches I • Typological Species Concept: species are a 'type' of organism…Plato • Existence of a limited number of universals • No regard to variation in a species • "Species are as many as were created in the beginning by the Infinite." (Linnaeus, 1758) Non-Evolutionary Approaches II • Essentialism • Similar to typological thinking • Species consist of individuals sharing the same essence • Species are separated by sharp discontinuity • Each species is constant through time • Limits to variation within a species Non-Evolutionary Approaches III • • • • • Nominalistic Only Individuals exist There are no “real” universals Popular in France in late 1800s Nature produces individuals and nothing more…no species. • Species are only mental constructs 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Evolutionarily Species Concepts • The lineage species concept • A single species forms an evolutionary lineage separated from other species. • They are not in genetic contact with other species • Therefore, the species concepts might be criteria for determining the split between species. 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Biological Species Concept • Species are groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups (Mayr, 1940). 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Problems with the BSC • What do you do with asexual organisms, and what do you do with organisms that occasionally form hybrids with one another? Other difficulties include: • What is meant by “potentially interbreeding • Fossils---for example, it is not really possible (or very meaningful!) to figure out whether a trilobite living 300 million years ago would have interbred with its ancestor living 310 million years ago. 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Evolutionary species concept • Evolutionary species concept: A species is a single lineage of ancestor-descendant populations which maintain its identity from other such lineages and which has it own evolutionary tendencies and historical fate (Simpson, 1961; Wiley, 1981). • Difficult to apply, so we use the following: 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Phylogenetic species concept • A species is the smallest diagnosable cluster of individual organisms within which there is a parental pattern of ancestry and descent (Cracraft 1983). 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Six Giraffe Species? Approximate geographic ranges, pelage patterns, and phylogenetic relationships between giraffe subspecies based on mtDNA sequences. (From: http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2254591/ ) https://www.wvup.edu/.../Biology%20461%20Species%20and%20Sp... Other concepts • Cohesion species concept: A species is the most inclusive group of organisms having the potential for genetic and/or demographic exchangeability. (Templeton, 1989) • Ecological species concept: A species is a set of organisms exploiting (or adapted to) a single niche (Ridley 1993). • Recognition species concept: A species is the most inclusive population of individual biparental organisms which share a common fertilization system. (as defined by Paterson, 1985; in Templeton, 1989). • Isolation species concept: Species are systems of populations: the gene exchange between these systems is limited or prevented by a reproductive isolating mechanism or perhaps by a combination of several such mechanisms. (as defined by Dobzhansky 1970; in Templeton, 1989) 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Speciation • Formation of new species (a) Morphospecies. Viewed today, at one moment in time, species A, C, and E are clearly distinct, demarcated by current natural discontinuities between them. (b) Paleospecies (chronospecies). Viewed historically, through time, discovered fossil intermediates (B and D) fill in the missing gaps above, giving us a more or less continuous series with no obvious discontinuities between them. https://www.wvup.edu/.../Biology%20461%20Species%20and%20Sp... Allopatric Speciation • The simplest and most common mechanism of speciation is allopatric speciation: 2 groups of one species are isolated geographically, and diverge into separate species. www.bios.niu.edu/johns/bios103/speciation.ppt Allopatric Speciation • Results from populations of a single species becoming separated from each other and evolving in different ways • 1. Geographic isolation • 2. Adapt to different environments • 3. Can no longer interbreed with each other Early fox population Northern population Spreads northward and southward and separates Arctic Fox Adapted to cold through heavier fur, short ears, short legs, short nose. White fur matches snow for camouflage. Different environmental conditions lead to different selective pressures and evolution into two different species. Southern population Gray Fox Adapted to heat through lightweight fur and long ears, legs, and nose, which give off more heat. Rocky Mountains may be a barrier that prevents Canada lynx from breeding with populations on the other side of the mountain. U.S. Fish & Wildlife Service Image PA NG AE A 120° 80° LAURASIA 40° 80° 120° 120° 80° 80° 120° GONDWANALAND 135 million years ago 225 million years ago Continental drift NORTH AMERICA EURASIA AFRICA 120° 80° SOUTH AMERICA INDIA MADAGASCAR 120° 120° 0° AUSTRALIA ANTARCTICA 65 million years ago Present 40° 120° Reproductive barriers Prevent gene flow Peripatric Speciation • • • An isolated group is severed from the original population Rare genes move to fixation If these genes are associated with reproduction, sexual selection takes over, then speciation has occurred as they may not mate with the original population 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Hawai’ian Drosophila 500 species!!! http://dbs.umt.edu/courses/biol223/documents/speciation2232006post.ppt. Parapatric speciation • In parapatric speciation there is no specific extrinsic barrier to gene flow. • The population is continuous, but nonetheless, the population does not mate randomly. • Mating with geographic neighbors more often. • In this mode, divergence may happen because of reduced gene flow within the population and varying selection pressures across the population’s range. 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Sympatric speciation • Does not require large-scale geographic distance to reduce gene flow between parts of a population. • Exploiting a new niche may automatically reduce gene flow with individuals exploiting the other niche. • This may occasionally happen when, for example, herbivorous insects try out a new host plant. 163.238.8.180/~fburbrink/Courses/.../Evolution%2017%20ppt.ppt Species and Speciation Sympatric Speciation Cichlid fish in Lake Barmobi Mbo in Cameroon, Africa—an isolated volcanic lake. Nine species, all more closely related to each other (by DNA evidence) than to similar fish in other lakes. Lake has no distinct geographical zones, and the fish can easily swim anywhere in it. They feed in different locations, but all breed in the same location, close to the bottom. An example of sympatric speciation, but the mechanism is not clear. www.bios.niu.edu/johns/bios103/speciation.ppt Sympatric Speciation Polyploidy July 27, 2005 Two Species Become One, Study Says The newly discovered Lonicera fly species arose through hybridization, a new study says. Two other fly species mated and formed a hybrid, a combined form that cannot mate with its fellow hybrids. Given a separate niche in which to evolve—in this case, an alien huneysuckle imported to the U.S.—a hybrid animal can become a full-fledged new species, according to researchers. Photograph by Guy Bush Ring Species http://dbs.umt.edu/courses/biol223/documents/speciation2232006post.ppt. Speciation & Macroevolution The Gradualism model suggests that change is gradual with the accumulation of unique morphological adaptation. - The Punctuated Equilibrium model suggests that rapid change occurs, with a new species “erupting” from the ancestral lineage and then staying the same thereafter. www.eeescience.utoledo.edu/.../Lecture%207%20-%20macroevoluti... Mesozoic Cenozoic Marsupials (kangaroos, etc.) Rabbits Rodents Primates Bats Insectivores Carnivores Whales Even-toed hoofed mammals Odd-toed hoofed mammals Elephants Monotremes (platypus, etc.) Extinction • • Background extinction – inability to adapt to changing environment Mass extinction – periodic large-scale extinctions resulting from asteroid impacts or other causes – Reshuffle life on the planet – Winners become losers and vice versa • E.g. 65 mya mammals take over from reptiles and underwent adaptive radiation Cenozoic Era Period Quaternary Millions of years ago Today Bar width represents relative Species and families experiencing number of living species mass extinction Extinction Tertiary 65 Extinction Mesozoic Cretaceous Jurassic 180 Extinction Cretaceous: up to 80% of ruling reptiles (dinosaurs); many marine species including many foraminiferans and mollusks. Triassic: 35% of animal families, including many reptiles and marine mollusks. Triassic 250 Extinction Permian Carboniferous Paleozoic Current extinction crisis caused by human activities. Many species are expected to become extinct within the next 50–100 years. 345 Permian: 90% of animal families, including over 95% of marine species; many trees, amphibians, most bryozoans and brachiopods, all trilobites. Extinction Devonian: 30% of animal families, including agnathan and placoderm fishes and many trilobites. Extinction Ordovician: 50% of animal families, including many trilobites Devonian Silurian Ordovician Cambrian 500