Biology 3.5 Patterns of Evolution Credits: 4 External ! Before we begin… From the examination specifications… Giraffe Evolution? Revision 12 bio stuff… Stuff u should know from Year 12 • Natural Selection – Directional – Disruptive – Stabilising • • • • • • • Founder effect Population bottlenecks Genetic drift Mutation Gene flow Migration Factors affecting allele frequencies… Darwin vs Lamarck Natural Selection in a NutSHELL Variation Individuals vary in phenotype (physical traits). Some variants are better suited to the current environmental conditions. The better suited survive better and leave more offspring. Natural Selection The alleles of the better suited increase in the population, they are said to be ‘selected for’. Inheritance Variations are passed on to offspring. If the selection pressure is maintained each new generation contains proportionally more descendants from individuals with favourable characteristics than those with unfavourable. Some variation is lost. Natural Selection Requires: Variation Selective Pressure Inheritance Result: Selection of the ‘fittest’ variants and their genes. Bio 3.5 Level 3 from here The “Species” Concept “a group of actually or potentially interbreeding natural populations that is reproductively isolated from other such groups” Main Problem: - Closely related species produce fertile offspring eg Canis spp. - Genetically isolated species may be morphologically similar = cryptic species (morph = “body”) Main Solution: - Use DNA analysis to clarify relationships between closely related species. - BUT: how much difference = different species, subspecies….?? Extinct: 10,000 years bp Clines & Ring Species • Cline: A gradation in one or more characteristics within a species esp. between different populations. • Ring Species: A series of neighbouring populations that can interbreed, but for which there exist at least two "end" populations in the series that are too distantly related to interbreed In this diagram, interbreeding populations are represented by coloured blocks. Variation along a cline may bend right around, forming a ring. Larus (gull) ring species No, gull away! Are you my type? A Herring Gull, Larus argentatus (front) and a Lesser Black-backed Gull. Larus fuscus (behind) The Larus gulls interbreed in a ring around the arctic (1 : Larus argentatus argentatus, 2: Larus fuscus sensu stricto, 3 : Larus fuscus heuglini, 4 : Larus argentatus birulai, 5 : Larus argentatus vegae, 6 : Larus argentatus smithsonianus, 7 : Larus argentatus argenteus Neighbouring groups can hybridise (breed together) but sufficient differences exist to prevent groups 1 & 7 breeding. Californian Salamander Ring Species The many subspecies of Ensatina salamanders in California exhibit subtle morphological and genetic differences all along their range. They all interbreed with their immediate neighbours with one exception: where the extreme ends of the range overlap in Southern California, E. klauberi and E. eschscholtzii do not interbreed. So where do we mark the point of speciation? Stages in Species Development #1 • Species rarely explode suddenly into existence (species formation is usually slow) • General pattern: – Homogenous population splits (cause = geographical barrier) – Different natural selection pressures, mutations gene frequencies change – Races form as gene flow reduces, factors preventing mating begin (“prezygotic”) – Gene flow further reduces, post zygotic factors occur (hybrid sterility – eg as in mule) – Now = two different species. Variation in Human Skin Colour Stages in Species Development #2 - A generally predictable series of events occurs as a homogenous ancestral population evolves into two separate species. - The key to this is build up of genetic differences as a population is split into two populations. Barriers to frequent mating mean that differing natural selection pressures and mutations are not shared populations go down different genetic pathways. - Eventually differences build to where the populations become separate races. - With enough differences races become difference subspecies, then separate species. Extinction Extinction Sumatran tiger Sumatran tiger: http://en.wikipedia.org/wiki/Sumatran_tiger Sumatran tiger clip: http://www.theawl.com/2011/05/get-a-good-look-at-these-awesome-tigers-theyre-almost-extinct Call of Life (extinction video trailer): http://www.calloflife.org/p-trailer.htm Extinction • A natural process – all species that have evolved will eventually go extinct • Duration of persistence of a species varies (often from 1 million years for complex organisms to 10-12 million years for simple organisms) • Extinction and mass extinction provides opportunities for other organisms to evolve and fill vacant ecological niches. Humans: How long have we been around? When will we become extinct? • Anatomical modernity: 200,000 years ago • Behavioural modernity: 50,000 years ago • Future…”currently using resources at rate of 8 planets worth” “Fred” a preserved dodo, Raphus cucullatus Haasts Eagle & moa Huia Coelacanth Latimeria chalumnae (60kg, 170cm long) Coelacanths were thought to have gone extinct in the Late Cretaceous (~65mya), but were rediscovered in 1938 off the coast of South Africa. 2 Known extant species. The coelacanth has been nicknamed a “living fossil”, because its fossils were found long before the actual discovery of a live specimen. The coelacanth is thought to have first evolved approximately 400 million years ago. Processes of Evolution The Role of Mutation • Notes + BZ • 90,92-98 The Role of Gene Flow • Notes + BZ • BLB 158-160 Oligocene Drowning • http://www.odt.co.nz/lifestyle/magazine/251 158/theory-flounders • http://www.stuff.co.nz/environment/215430/ NZ-was-never-underwater-scientists Kiwi an Australian? • http://www.nzherald.co.nz/nz/news/article.cf m?c_id=1&objectid=11259906 • http://www.adelaide.edu.au/news/news7068 2.html Speciation • “Process by which one species gives rise to two or more new species” • Multiplication of spp. not gradual change over time Allopatric Speciation Usually: • Pops get geographically separated (eg by river) • Gene flow reduces, Genetic isolation occurs. • Diffs in natural selection can cause diffs in allele frequencies between the pops over time. AND: mutations occurring in one population will not be transferred by migration to other populations (no gene flow) • Diffs accumulate, when the pops are reunited they may now no longer interbreed = separate spp. • NZs isolation/islands has led to many egs of allopatric speciation. (changing sea levels, mountain building, ice ages have also had an impact) Allopatric Speciation Allopatric Speciation: Snapping Shrimp Example 15 different species of shrimp on EACH side of the Isthmus of Panama (3mya). How could this occur? In a single population of shrimp (before the isthmus), a mutation that arose in one individual could eventually spread through the whole population as the shrimp mated with each other. But once there is a barrier splitting the population in half, a new mutation can only spread through half the population. That is why a lack of interbreeding means the two populations evolve separately. There can also be slightly different factors for survival in the different areas: maybe the temperature or currents are different on each side; maybe the food sources are different. This can also help the populations diverge, or become different, from each other. Reptiles again… Allopatric Speciation Questions 1. Why have many NZ birds lost the ability to fly (cf to their Aussie relatives)? 2. Glaciation creates many isolated mountaintops – how would this contribute to allopatric speciation? 3. How could sea level rise/fall create new species through allopatric speciation? 4. Describe how an ancestral robin species gave rise to the Chatham Island robin and the mainland robin. 5. Describe ALL potential geographical barriers (read the sheet “geographical barriers” 6. Explain how one NZ plant and one NZ animal came about as a result of allopatric speciation (read the sheet “allopatric speciation in NZ animals and plants” Sympatric Speciation • ‘speciation NOT involving a period of geographic separation” • Much rarer than allopatric speciation • More common in plants • Rest of notes on OHT (sigh) Nonallopatric Speciation occurs with no evidence of physical barriers Geography is not the only way a population can separate The huge variety of cichlid fishes in African lakes are found nowhere else; yet the lakes are evolutionarily young and without barriers. Individuals can speciate while living in different components of the environment. African cichlid fishes show very different feeding specializations Polyploidy Non-disjunction • BZ figs pg 114 • Animation • Inc polyploid def This image shows haploid (single), diploid (double), triploid (triple), and tetraploid (quadruple) sets of chromosomes. Triploid and tetraploid chromosomes are examples of polyploidy. Inducing Non-Disjunction • Done deliberately to create new plant varieties • Often larger, more vigorous than parents • Uses seeds/seedlings soaked in colchicine / N2O* gas – Inhibits spindle fibre formation – Chromosomes fail to separate – Resulting gametes may lead to polyploid plants • Propagated asexually or crossed if fertile *yes that is nos, nitrous oxide…. Polyploidy as a Source of Variation #1 • Background • Allo & auto • Hybrid vigour… Polyploidy in Manuka Don’t get confused • • • • • Polyploidy Polysomy Anueploidy? Allopolyploidy Autopolyploidy Teosinte Modern Corn (~1000yrs) Wheat Spelt • Where does this fit in wheat development? • Hybrid of emmer wheat • & goat grass in near east before the hexaploid bread wheat appears • See footnotes for more… • Nb bread wheat 8000 years ago • http://en.wikipedia.org/wiki/Spelt Reproductive Isolating Mechanisms • Notes uploaded to weebly… Lacewing Songs: • http://www.pbs.org/wgbh/evolution/library/0 5/2/swf_pop/l_052_01.html Evolutionary Relationships - Terms • Phylogenetics: the study of evolutionary relatedness between groups of organisms. Relatedness is determined by DNA sequencing data and comparing morphological data • Phylogeny: The evolutionary development and history of a species or higher taxonomic grouping of organisms. • Cladogram: Diagram which shows ancestral relations between organisms • Cladistics: method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants (and nothing else). Cladograms • Show ancestral relations between taxa – Using DNA analysis or morphological comparisons • Species are at the “leaves” • Common ancestor at the “trunk” • Have an implicit time axis (runs forward from base to leaves) but: problems of scale, data quantity & quality • May show extinct species, but: DNA from extinct species is rare canids are an old lineage, separating from the Simplified Canid Phylogeny *The other carnivores about 60 million years ago. of a "wolf" branch, a "South American" (shown in the cladogram below)Separation branch, and a "red fox" branch occurred more time recently, 7-10 million years ago. a clade *Mitochondrial DNA analysis of both modern and historical specimens of red wolves failed to distinguish red them as a species separate from gray wolves or coyotes. They appear to be a hybrid species, and can interbreed with either gray wolves or coyotes. *Two different dates for the origin of dogs have been suggested. Mitochondrial DNA analysis suggests a date between 60-100,000 years ago -well before the beginning of human agriculture. Other genetic and archeological evidence suggests a more recent date -- about 15,000 years ago. Neolithic cave drawings also show dogs hunting with humans. common ancestor species *All domestic dogs are the descendants of a few ancestral wolf stocks originating in Asia. Surprisingly this includes New World dogs, who were once thought to have been independently domesticated from New World wolves. Molecular Phylogeny (DNA analysis) may revise past phylogenies (based on morphology)… The Squamata, or the scaled reptiles, are the largest recent order of reptiles, comprising all lizards and snakes. Hedges, S. Blair, and Poling, Laura L. A Molecular Phylogeny of Reptiles. Science, Vol. 283, No.5404, pp.998-1001 • The study also cast in doubt the relationship between the tuatara and squamates. While fewer gene sequences were available for the tuatara, six of eight comparisons showed closer affinities with archosaurs or turtles, while only two showed squamates as the closest relative. While the results of this study are not conclusive, it clearly demonstrates that we don't know all that we thought we knew about the phylogenetic relationships of living or fossil reptiles. http://home.pcisys.net/~dlblanc/articles/TurtleP hylogeny.php Convergent Evolution • The evolution of the same biological trait in unrelated groups / species. • Examples: – Shark, icthyosaur, dolphin, penguin (a fish, reptile, mammal and bird respectively) are unrelated but have evolved a similar streamlined shape and “fins” in response to their environment (water) which provides a common selection pressure (in this case fast movement through water) – Unrelated plants have evolved water storage tissue (succulent tissues) eg Euphorbia, cacti in response to their environment (dry desert) which provides a common selection pressure (need to store water during prolonged dryness) • Analogous structures “structures that are alike in function but have a different evolutionary origin” – Egs: wings of insects and birds; mammalian and octopus eye Human eye vs Octopus eye Unrelated organisms (vertebrate vs mollusc) Developed similar eye as a result of selection pressure for well developed sight (see prey clearly, 3D) The eye is an analogous structure Dolphin Icthyosaur (2-4m) Penguin Shark Convergence in Plants Ferocactus pilosus (Mexican lime cactus) These unrelated plants have separately evolved the ability to store water in their stems. This is a response to the natural selection pressure of dryness in the desert. The swollen stems are an example of an analogous structure Example (reading) • http://blogs.discovermagazine.com/notrocket science/2012/11/20/the-deadliest-sea-snakeis-actually-two-look-alike-species/#more-7978 • http://tinyurl.com/SSSSSSSSNAKE Divergent Evolution • When one ancestral group evolves into two or more species, usually in different habitats • Features: – Accounts for most evolutionary change – Often due to ancestral spp. Increasing range / colonising new areas / habitats (new ecological niches) The different conditions cause different selection pressure different genetic pathways genetic isolation speciation • Alternatively: – Sequential evolution: small changes build up over time until a new species emerges (aka anagenesis, phyletic graduation) – Budding: a new species branches off while the ancestral species remain unchanged. • Cladogenesis: When a whole new group of organisms evolves (eg primates) Homologous Structures “The structures shared by a set of related species because they have been inherited, with or without modification, from their common ancestor” For example, the bones that support a bat's wing are similar in structure, type and number to those of a human arm. Differences are caused by differing natural selection pressures. Eg selection for traits enabling flight favours thin light bones in the bat. What pressures result in the other limbs pictured? Homologous Structures • http://collections.tepapa.govt.nz/exhibitions/ Whales/Segment.aspx?irn=159 Adaptive Radiation • “The diversification of a group of organisms into species filling different ecological niches”. • Can occur very rapidly, usually when a large number of ecological niches are vacant. • Example par excellence: – Dinosaur extinction 68mya opened up many niches for exploitation (eg Brontosaurus death opened up a large browsing herbivore niche). Relatively non specialised mammals (eg Megazostrodon were, as adaptable ‘generalists’, able to fill these niches quickly and through natural selection speciate into new forms. • Note: In a discussion you MUST involve natural selection AND niche. – Eg for above example: variation in ancestral mammal species size diet some bigger & better able to use plants survived better & reproduced more, passed those genes on further selection for various traits elephant! Asteroid: “Bye bye dinos, vacate your ecological niches!” Mammals: “Thanks a lot for the niches, we will fill those!” Megazostrodon model, Natural History Museum, London • Other examples: – Galapagos Finches: 1 South American finch evolved into 14 spp. occupying different niches (desert, grassland…) on the Galapagos Islands • NZ Examples: – 100 spp. of Hebe plants (specialised into coastal, forest, alpine… niches) – 10 spp. of Powelliphanta snails (+ subspecies, over different niches) – NZ parrots (kakapo, kea, kaka) from one ancestor 100mya (forest vs alpine) • Note: In some of these egs radiation was very fast (many vacant niches) and involved the founder effect. Powelliphanta spp. Ale – see folder for text Weird Mammal Groups #1 • Ungulates (meaning roughly "being hoofed" or "hoofed animal") a – The odd-toed ungulates are browsing and grazing mammals, such as horses, tapirs and rhinoceroses, whose hooves each feature an odd number of toes – Even toed ungulates: This group includes pigs, peccaries, hippopotamuses, camels, chevrotains (mouse deer), deer, giraffes, pronghorn, antelopes, sheep, goats, and cattle. Weird Mammal Groups #2 • Placentals – Eg dog, horse • Marsupials – Eg wallaby • Monotremes – Eg platypus Hawaiian Honeycreepers Hawaiian islands: volcanic origin, variety of habitats. In absence of other bird spp. they radiated to fill numerous niches Galapagos Finch Niches http://web.visionlearning.com/cus tom/biology/animations/darwin_f Anims… inches_working.shtml http://faculty.massasoit.mass.edu/whanna/1 22/page4/page7/page58/page58.html http://faculty.massasoit.mass.edu/whanna/1 22/page4/page7/page58/page58.html Co-Evolution • Change in the genetic composition one species (or group) in response to the change in another. • Often occurs when close ecological interaction (symbiosis) occurs. Eg: – Between parasite & host - exploitation – With flower shape & pollinator - mutualism – Between predator & prey – exploitation • Examples: – Lions: speed, strength, co-operative hunting (to catch gazelle) – Gazelle: speed, size, strength, horns, darting behaviour (to escape lion) • An “evolutionary arms race” one species evolves to respond to the other. • Each party exerts selection pressures on the other over time the spp may become mutually dependent on each other. • Relationship may become so close that extinction of one app means extinction of the other – Pollination Syndromes: where only one animal can pollinate only one specific plant (often the pollinator beak shape co-evolves with the flower shape) – Eg Adams’ mistletoe probably relied only on one NZ bird for pollination (pollinator not yet confirmed) when this birds dropped in numbers Adams mistletoe became extinct. Adams Mistletoe A victim of pollination syndrome? Co-evolution example Pseudomyrmex ant collecting protein-rich Beltian bodies from a bullhorn acacia, Costa Rica. This is one of the most famous mutualisms of all, the relationship between Pseudomyrmex ants and Acacia trees. The ants defend these small trees against herbivorous insects and vertebrates. The ants also chew away and sting any encroaching plants, clearing an area that may be up to 4 yd (4 m) in radius. In return, the plants give the ants food, such as the yellow Beltian bodies seen here, and nectar from extra-floral nectaries. The Beltian bodies contain proteins and lipids and are produced on the youngest and most delicate leaves. The plants also produce thorns that the ants hollow out for nests. Q. How could this evolve? Anna's Hummingbird (Calypte anna) and bottlebrush sp. The needle like beak and hovering ability of a hummingbird is allows it to extract nectar (and pollinate) funnel shaped flowers or flowers with no “landing pads” Dactylanthus taylorii Pollinated by native bat with which it shows coevolution. Bat: nocturnal, blind BUT good sense of smell Flowers: drab, open at night, strong scent (like a bat), wide shape (for easy access) Punctuated Equilibrium • Evolutionary model where there are long periods of little change in a spp punctuated by short bursts of rapid change. – Long periods of no evolutionary change (stasis) – Stasis punctuated by short periods of evolution producing new species rapidly – Stimulus for evolution = environmental change – Species’ spend most of existence in stasis – If Correct: no transitional fossils, sudden appearance of new types Gradualism • Evolutionary model where the accumulation of changes resulting in speciation occurs slowly and steadily – Evolution proceeds slowly but continuously in response to selection pressures – Eventually changes in adaptive characteristics accumulate until speciation occurs – If correct transitional forms should be seen (as is the case with horse evolution) – Example: Trilobites changed gradually over three million years A Trilobite fossil: Kainops invius Trilobites… • Hard-shelled, segmented creatures lived over 520 million years ago in Earth's ancient seas. • Extinct before dinosaurs arrived • Key creatures of the Paleozoic Era, (1st era with complex life) • Fossils found in rocks in all continents Body plan: 3 main parts – head, segmented thorax, and a pygidium (tail piece) Trilobite means three lobed (see right). Huge morphological diversity in trilobites but all have this basic structure Size: 3mm – 300mm www.trilobites.info Evidence for evolution • http://www.sumanasinc.com/webcontent/ani mations/content/evolution/evolution.html Poster Task Rest of the period to: • Present an A3 colour poster on an example of convergent evolution, allopatric speciation or sympatric speciation. • You must: • • • • Define the type of evolution speciation Give a captioned example (no kea, kaka, butterfly) Explain how this example came about Name it! Evidence for Evolution 1) Fossil Record Fossil = preserved impression turned into rock or mineralised remains of past organisms Fossil record Sedimentary rock forms in layers which may contain fossils. Older strata/fossils usually found deeper (exception = after uplift). This provides a record of: Appearance/disappearance of species Diversity at various times Evidence of change of ancestral forms to modern forms Fossil age can be determined by: dating rock layers, comparing to fossils in same strata elsewhere. Problems: Destruction of fossils (by subduction), some organisms don’t fossilise well (eg soft bodied animals), most fossils are still buried. Archaeopteryx represents a transitional species between reptiles and birds Fossil Record 2) Comparative Anatomy “Comparison of body structures between different species” Homologous Structures • Similarity of structure between related species indicates evolution from a common ancestor. Eg inheritance of pentadactyl limb in different mammals Vestigial Organs • Organs reduced in function size but remnant is shared with common ancestor or closely related species. Eg human appendix = reduced caecum, lost function of digesting cellulose, whale pelvis remnant - now no hind limbs. How would natural selection against these structures operate? Whale What vestigial organs do kiwi have? Snake 3) Biogeography “The study of geographical distribution of species” Species on a given island may more closely resemble species on a nearby mainland rather than species on a distant island (even though the habitats may be similar). Eg Galapagos finches similar to mainland S.America finches, not similar to birds 1000s km away on Cape Verde islands. Adaptive radiation, allopatric speciation important here. Absence of certain taxa can be explained by their evolution after a separation event. Eg placental mammals evolved after NZ split from Australia 85mya; this explains their absence in NZ. BUT we have examples of old Gondwanan lineages eg tuatara, frogs. Fossils of related organisms found on separate continents can be explained by continental drift. Eg glossopteris (a fern) evolved in Gondwana (350-230mya). Gondwana split up into modern day Antarctica, Australia, India, Africa, S. America and fossils of it are found in all locations. It couldn’t possibly have evolved after the split up and spread to each location. Therapods evolved in one place 140mya, spread out, continents split, therapods left fossil remains of related species in spread apart locations. Much more plausible than a recent independent origin of each species in each location. Biogeography Some organisms that evolved in Gondwana have left fossils spread out amongst modern day continents. Unlikely that they could have evolved recently and spread out over the large distances involved. Eg how could the land reptile lystrosaurus get from India to Antarctica? 4) Molecular Biology • Genetic Code: this is the base sequence of DNA. Organisms which have a recent common ancestor will share more of the base sequence than those that are unrelated. If the mutation rate is known then base sequence difference between two species can be extrapolated to determine when the last common ancestor was; • Proteins: more closely related species have more similarity in their proteins (this reflects a similarity in the underlying DNA that codes for the proteins). • Hox Genes: a group of related genes that control structure and orientation of organisms – critical for development and placing of body parts. The homeobox is a sequence within a hox gene that makes a protein that acts as a switch for the gene. The homeobox has been found to be highly conservative (changes little) across evolutionary lineages. A fly functions perfectly well with a chicken hox gene in place of its own. Interpretation: The human/chimp lineage split relatively recently – there has not been enough time for mutations to cause differences in the bases of the cytochrome c gene so the amino acid sequence is identical The human/yeast lineage split a relatively long time ago, there has been a lot of time for base differences to accumulate… Rate of DNA hybridisation Via the process of DNA hybridization, scientists can accurately determine the degree of relatedness between various groups of species. Matches in base sequence between species indicate a high degree of relatedness. For example, in the figure on the right you can see that more matches are made between a human and a chimp than between a human and a chicken. This indicates that the human and the chimp shared a more recent common ancestor than the human and the chicken and are closely related. Don’t confuse the two!