Topic 3: Systematics II Schools of Systematics What are the schools of thought of systematics? How does one make a cladogram? What are higher taxa & ranks? • • If the system is based on evolution, then it is phylogenetic How should they be used in this course? • These underlying principles are typically evolution and parsimony Applying phylogenies – Evolution and homoplasy in Anolis Schools of Systematics Systematics – The clustering of groups or organisms based on a unifying set of principles (evolutionary, or otherwise) Evolutionary Systematics Phenetics Cladistics Model-based approaches ______________ ______________ ______________ ______________ What is Evolutionary Systematics? Oldest approach to reconstructing phylogeny __________________ Often ______ objective – “Anything goes” What is Evolutionary Systematics? An evolutionary tree Note: ancestors, time, and anagenesis Ancestors occur and time is recognized Anagenesis is recognized Both monophyletic and ______________ groups are recognized No longer used Basal Archosaur Time What is Evolutionary Systematics? Clustering is based on both _____________ and ________________ features Evolutionary trees are produced No explicit procedure to reconstructing phylogeny The scientist’s opinions about character weighting and relationships are important Basal Diapsid Basal Sauropsid Basal Amniote 1 What is Phenetics? A response to the lack of objectivity of Evolutionary Systematics Not necessarily __________________ What is Phenetics? By looking at gross similarity, the analysis can be confounded by _______________ and __________________ Modern uses: ‘True’ evolutionary relationships are not knowable Measures overall ______________ among taxa Low taxonomic levels Morphometrics and some immunological data Does not differentiate between ancestral and derived (characters are not polarized) Explicit quantitative procedure What is Cladistics? Also called Phylogenetic Systematics What is Phenetics? A ______________________ is produced Note confounding effects of anagenesis and convergence Evolutionary Tree Originated in the 1950s by Hennig, popularized in the 1960s, further developed after that Response to the need for a ______________ system that was ________________ and _________________ Therefore, phylogenetic Phenogram What is Cladistics? Also called Phylogenetic Systematics Only ____________________ are used to cluster taxa hierarchically __________________, not anagenesis, is the focus of cladistics Premise that little is known about character evolution What is Cladistics? Also called Phylogenetic Systematics Used extensively from the 1960s to present day Used to reconstruct phylogeny and study character evolution Well implemented in computer algorithms Not explicitly model-based Parsimony is the guiding principle 2 Cladistics: How to make a cladogram _______________ Timeless and without ancestors A representation of character state distribution An hypothesis of relationship _______ an evolutionary tree (OG (A (B (C, D)))) Cladistics: How to make a cladogram A B C D E How do these “Caminacules” differ from one another? Cladistics: How to make a cladogram A B C D E Code each of the characters (columns) in a table (taxa in rows) 1. Single or close together hind limbs (0), OR 2 separate hind limbs (1) 2. Front limbs are short (0), OR lengthened (1) 3. Abdomen is unpatterned (0), OR patterned (1) 4. Abdomen is flat (0), OR bulbous (1) 5. Two (0) front limbs, OR four (1) 6. One or no eyes (0), OR two laterally-directed eyes (1) 1 Cladistics: How to make a cladogram Taxon A 1 2 3 4 5 6 B C D E Cladistics: How to make a cladogram A B C D E 1 What is the likelihood approach? Cladistics: How to make a cladogram However, with many taxa, finding the best topology is computationally intensive With m taxa, then number of rooted topologies, n, is: n= (2m − 3)! m− 2 2 (m − 2)! Developed in the 1960s to present Places phylogenetic reconstruction in an explicitly ____________________ framework Realization that we do know things about character evolution, particularly DNA Find the tree with the greatest likelihood: If m = 5, then n = 7!/(8 x 6) = 105 L ∝ P( Data | Tree) Maximum parsimony __________ ___________________________ ___________________________ What is Bayesian Inference? Developed in 1764 by Thomas Bayes Applied to phylogenetics in 1996 Builds on likelihood approaches by also allowing for the incorporation of ___________________ Allows calculation of the actual probability of a tree: L P( data | tree) P(tree) P (tree | data ) = P(data ) Why are model-based approaches so powerful? Both likelihood and Bayesian approaches are computationally intensive Allow us to use knowledge of character evolution in reconstructing phylogeny, which parsimony doesn’t really do Primarily applied to DNA sequence data, but recently, even morphological data can by used Why are model-based approaches so powerful? What do we know about DNA evolution? (More in other classes) 1 What are higher taxa and ranks? What are higher taxa and ranks? What do they mean? Domain Kingdom Phylum Class Order Family Genus • Erected to recognize diversity in Archae, Eubacteria, Eukarya • Major divisions within Eukarya (Plantae, Animalia, Fungi, etc.) • Distinct body plans within Animalia (Plantae and Fungi use Divisions) Domain Kingdom Phylum Class Order Family Genus • Some sub-groupings Species Species Subspecies Subspecies • ____________________________________ • Some distinct group within a species? Ranks What do they mean? Are they real? Are they comparable? How should they be used? Hierarchical framework of classification What are higher taxa and ranks? • 3 domains are monophyletic, no evidence that a Domain is something biologically real • Most are monophyletic, no evidence of biological reality • Recent evidence suggests that evolution of developmental genetic “kernels” defined __________ before the Cambrian explosion* Domain Kingdom Phylum Class Order • _________________ of biological reality Family Genus Species Subspecies • Most scientists agree that species are actual ___________________________ • Little evidence for biological reality * Davidson & Erwin. 2006. Science 311:796. What are higher taxa and ranks? Domain Kingdom Phylum Class Order Family Genus Species Subspecies What are higher taxa and ranks? Are they real? How should they be used? Most don’t have a specific meaning, are not biological entities, and aren’t comparable to other groups of the same rank Trend to not use ranks Domain Kingdom Phylum Class Order Family Genus Species Subspecies Are they comparable? Families of Urodela (Salamanders): Fig. 3-1: Pough et al. 2004. An Example – Evolution & Homoplasy in Anolis • Approximately 400 species of Anolis, ~125 of which occur in the Caribbean Including this course! Use monophyletic groups (clades) to refer to higher taxa _________________ are obviously useful, maybe _____________ as well Photos © AS Munoz, G White, J Hance, PJ Bergmann 1 An Example – Evolution & Homoplasy in Anolis Anolis Ecomorphs • Many species occur in the Caribbean • On the Greater Antilles, “___________” have evolved Twig giant • The same ______________, with similar morphology and ecology, have evolved on each of the four big islands Crown giant © J. Losos Trunk-crown Trunk-crown dwarf © Auburn Herp Soc Cuba Haiti/Dominican Rep. Trunk-ground Jamaica Puerto Rico Powell and Russell, 1992. Homoplasy in Anolis – different phylogenetic signal from different datasets © Reptilien-Center.de Replicate Evolution in Anolis Locomotor Morphology mtDNA Sequences Anolis ecomorphs evolved in a ______________ ______________ on each of the Greater Antilles From Jackman et al., 2000. From Jackman et al., 2000. 2
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