Systematics - BIO 615
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Systematics - BIO 615 Outline - History and introduction to phylogenetic inference 1. Pre Lamarck, Pre Darwin “Classification without phylogeny” 2. Lamarck & Darwin to Hennig (et al.) “Classification with phylogeny but without a reproducible method” 3. Hennig (et al.) to today “Classification with phylogeny & a reproducible method” Biosystematics History alpha phylogenetics Aristotle - Scala Naturae - ladder of perfection with humans taxonomy at top - DIFFICULT mental concept to dislodge! (use of terms like “higher” and “lower” for organisms persist) character identification evolution Linnaeus - perpetuated the ladder-like view of life linear, pre evolution descriptions phylogeny 1758 - Linnaeus grouped all animals into 6 higher taxa: 1. Mammals ( top ) 2. Birds collections classification biogeography 3. Reptiles 4. Fishes 5. Insects Describing taxa = assigning names to groups (populations) 6. Worms ( bottom ) = classification Outline - History and introduction to History phylogenetic inference Lamarck - 1800 - Major impact on Biology: 1. Pre Lamarck, Pre Darwin - First public account of evolution - proposed that modern species had descended from common ancestors over “Classification without phylogeny” immense periods of time - Radical! evolution = descent with modification 2. Lamarck & Darwin to Hennig (et al.) - Began with a ladder-like description… but considered “Classification with phylogeny but Linnaeus’s “worms” to be a chaotic “wastebucket” without a reproducible method” taxon 3. Hennig (et al.) to today - He raided the worm group to liberate disparate taxa “Classification with phylogeny & a - Early and MAJOR example of using data interpreted reproducible method” within an evolutionary framework to classify 1 Systematics - BIO 615 History Lamarck, J. B. 1809. Philosophie Zoologique History Perhaps the first branching diagram, “phylogentic tree” - Phylogenetics began before “evolutionary trees” published in the history of Biology - 1809 - Classifications were taken as proxies - representations of the “natural order,” these were the “trees” Lamarck’s ideas: - By putting taxa together in a group, one was saying Two origins of life (since both “these taxa are closer to one another than to were “equally primitive”) anything else - they belong together” Insects et al - side branch - “Closer” meant different things to different workers: e.g. Greater body form similarity / complexity Removed birds to side branch e.g. Greater evolutionary relationship rather than “below” mammals e.g. more shared homologies… (natural vs artificial) Lamarck, J. B. 1809. Philosophie Zoologique Biosystematics History alpha phylogenetics “Dans sa production des differents animaux, la taxonomy nature n'a pas execute une serie unique et character simple.” identification evolution (In its production of the different animals, descriptions phylogeny nature has not fashioned a single and simple series.) 1809 collections classification biogeography - Lamarck 1815 Phylogenetics began with the emergence of the idea of = the beginning of the end of the Scala evolution applied to classification Naturae 2 Systematics - BIO 615 History History Charles Darwin (& Alfred Russel Wallace) - “In regard to classification and all the endless disputes about the "Natural System," which no two authors define provided the mechanism (natural selection) to in the same way, I believe it ought, in accordance to my explain evolution - 1858 & 1859 heterodox notions, to be simply genealogical. Lamarck’s ideas, which lacked an accurate But as we have no written pedigrees you will, perhaps, say this will not help much; but I think it ultimately will, whenever mechanism, hadn’t spread heterodoxy becomes orthodoxy, for it will clear away an immense amount of rubbish about the value of characters, Darwin’s ideas did… and will make the difference between analogy and homology clear. The time will come, I believe, though I shall not live to see it, when we shall have very fairly true genealogical (That evolution occurred was apparent to many trees of each great kingdom of Nature.” who knew the fossil record, the problem, was “How?”) Darwin in a letter to Huxley, 1857 History History Classifications (starting in the mid to late 1800s) - Based on inferred evolutionary history - End of artificial classifications for convenience or for understanding the creator - But, “How does one infer evolutionary history?” no reproducible method, yet, to do so… Darwin (1859) On the Origin of Species History History Ernst Haeckel (late 1800s) The “method” of phylogenetic inference - Heavy supporter of (from which one may or may not derive a evolution, but not Darwin classification) that developed is referred to - “Ontogeny recapitulates as phylogeny” - not a law, as he thought Evolutionary Taxonomy - Coined terms: Ecology - spanned from the 1800s to the 1960s Phylum Phylogeny (1866) monophyletic - formed the “roots” of the discipline of polyphyletic phylogenetics 3 Systematics - BIO 615 History History Evolutionary Taxonomy (<1960s) Evolutionary Taxonomy (<1960s) - “Method” involved a reliance on - During the modern synthesis authority, not reproducible: systematics was marginalized “not a science” - Leading figures: Mayr & Simpson 1940s 1. Spend a lifetime learning as much as - Without a reproducible method, possible about the biology of a group hypotheses couldn’t be tested (morphology, development, fossils, etc.) 2. Publish a drawing of a phylogeny - No longer used, but exists in some forms based on one’s (informed) opinion today (in contrast to systematists who know nothing about their organisms!) = “Guessograms” Haeckel, 1872 - tree depicts 1) relationships, 2) timing of Haeckel, 1872 - tree lacks: 1) data? Characters supporting branching events, 3) diversity of groups through time branches? 2) quantification of support - strength of argument History Summary of contributions Evolutionary Taxonomy (<1960s) Good: “Know your organisms” Bad: not reproducible; no objective method; arbitrary, intuitive, no way to ET - many hypotheses generated, but no method to test. resolve conflicts Modern methods to test show many of the relationships to be “good”, or at least close… but many also rejected 4 Systematics - BIO 615 Outline - History and introduction to Lack of a reproducible method resulted in three phylogenetic inference major approaches: (Explosion in 1960s) 1. Pre Lamarck, Pre Darwin 1. Phenetics - similarity / distances only, not “Classification without phylogeny” evolution, not phylogeny, no weighting 2. Lamarck & Darwin to Hennig (et al.) 2. Cladistics - phylogeny inferred using “Classification with phylogeny but characters & parsimony without a reproducible method” 3. Hennig (et al.) to today 3. Statistical Phylogenetics - phylogeny “Classification with phylogeny & a inferred using corrected data & “best fitting reproducible method” model” History Phenetics - similarity only, not evolution, not phylogeny (1950-1960) - computerized, reproducible, “objective,” modern, sexy… - goal to remove all subjectivity, measure everything, use matrix of distances - sometimes called “Numerical Taxonomy” after book by Sokal & Sneath (1963) - cluster based on overall similarity - not evolutionary trees, but phenograms 1957 - first numerical phylogeny published (Michener & Sokal) History Phenetics - Problems Similarity sometimes reflects evolution… 1. Reproducible but no agreement on which clustering algorithm or statistic to use… Rhino Beetle Butterfly - different algorithms = different trees e.g. UPGMA, PCA… etc. True tree: - although there is only one true evolutionary tree, there are many alternative Butterfly Butterfly ‘similarity’ trees depending on the data & algorithm used Beetle Beetle 2. Similarity doesn’t always = evolutionary Rhino Rhino relationship! Phenogram Cladogram 5 Systematics - BIO 615 History And sometimes doesn’t… Phenetics - the good 1. Demanded explicit character analysis & laid Lizard Crocodile Bird groundwork for numerical phylogenetics True tree: 2. Still used for lower-level problems e.g. morphometrics, species demarcations… 3. And some higher-level problems Lizard Lizard Crocodile Crocodile Survives in various forms - an issue of debate… (see lecture on Distance methods) Bird Bird Phenogram Cladogram Definition - Phenetics Lack of a reproducible method resulted in three major approaches: Phenetics as originally applied, by Cain and Harrison (1960), refers to the "arrangement by overall 1. Phenetics - similarity / distances only, not similarity, based on all available characters without evolution, not phylogeny any weighting ...since it employs all observable characters (including of course genetic data when available)". 2. Cladistics - phylogeny inferred using characters & parsimony Cain, A. J., Harrison, G.A. (1960) Phyletic weighting. Proceedings of the Zoological Society of London 3. Statistical Phylogenetics - phylogeny 131: 1-31 inferred using corrected data & “best fitting model” History History Cladistics Cladistics 1950 - Willi Hennig “Grundzüge einer Theorie der - method explicit, reproducible phylogenetischen Systematik” - hypotheses of phylogeny could now be “tested” - German entomologist of Diptera - Hennig’s original method: - 1966 English translation of Phylogenetic 1. Distinguish homologies from analogies Systematics 2. Distinguish derived homologies - key: distinguish derived from (apomorphies) from ancestral ancestral homology (plesiomorphies) homologies - yields phylogenetic tree 3. Tree then built from apomorphies (evidence of common ancestry) - Vitriolic attacks