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Topic 3: Systematics II Schools of Systematics

 What are the schools of thought of systematics? • Systematics – The clustering of groups or based on a unifying set of principles (evolutionary, or otherwise)  How does one make a ?

• If the system is based on , then it is  What are higher taxa & ranks? phylogenetic  How should they be used in this course? • These underlying principles are typically  Applying phylogenies – Evolution and evolution and parsimony homoplasy in Anolis

Schools of Systematics What is Evolutionary Systematics?

 Evolutionary  ______ Oldest approach to reconstructing phylogeny Systematics

  ______ ______  ______

 Model-based  ______ Often ______objective – “Anything goes” approaches  No explicit procedure to reconstructing phylogeny  The scientist’s opinions about character weighting and relationships are important

What is Evolutionary Systematics? What is Evolutionary Systematics?

 An evolutionary tree  Clustering is based on both ______and  Note: ancestors, time, and ______features  Evolutionary trees are produced  Ancestors occur and time is recognized

 Anagenesis is recognized Archosaur

 Both monophyletic and ______groups Basal Diapsid are recognized Time  No longer used Basal Sauropsid

Basal Amniote

1 What is Phenetics? What is Phenetics?

 A response to the lack of objectivity of  By looking at gross similarity, the analysis can be Evolutionary Systematics confounded by ______and ______

 Not necessarily ______ Modern uses:  ‘True’ evolutionary relationships are not knowable  Low taxonomic levels  Measures overall ______among taxa  Morphometrics and some immunological data

 Does not differentiate between ancestral and derived (characters are not polarized)

 Explicit quantitative procedure

What is Cladistics? What is Phenetics? Also called Phylogenetic Systematics

 A ______is produced   Note confounding effects of anagenesis and Originated in the 1950s by Hennig, convergence popularized in the 1960s, further developed after that

 Response to the need for a ______system that was ______and ______

 Therefore, phylogenetic

Evolutionary Tree Phenogram

What is Cladistics? What is Cladistics? Also called Phylogenetic Systematics Also called Phylogenetic Systematics

 Only ______are used  Used extensively from the 1960s to to cluster taxa hierarchically present day  ______, not anagenesis,  Used to reconstruct phylogeny and is the focus of cladistics study character evolution  Premise that little is known about  Well implemented in computer character evolution algorithms  Not explicitly model-based  Parsimony is the guiding principle

2 Cladistics: How to make a cladogram

 ______ Timeless and without (OG (A (B (C, D)))) ancestors  A representation of character state distribution  An hypothesis of relationship  ______an evolutionary tree

Cladistics: How to make a cladogram

ABCD E

How do these “Caminacules” differ from one another?

Cladistics: How to make a cladogram

ABCD 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 1 2 3 4 5 6 A

B

C

D

E

Cladistics: How to make a cladogram

ABCD E

1 Cladistics: How to make a cladogram What is the likelihood approach?

 However, with many taxa, finding the best topology is computationally intensive  Developed in the 1960s to present  Places phylogenetic reconstruction in an

 With m taxa, then number of rooted explicitly ______framework topologies, n, is:  Realization that we do know things about 2( m − 3)! character evolution, particularly DNA n = 2m−2 (m − 2)!  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? Why are model-based approaches so powerful?

 Both likelihood and Bayesian approaches are  Developed in 1764 by Thomas Bayes computationally intensive  Applied to in 1996

 Builds on likelihood approaches by also allowing  Allow us to use knowledge of character evolution in for the incorporation of ______reconstructing phylogeny, which parsimony doesn’t  Allows calculation of the actual probability of a really do tree:  Primarily applied to DNA sequence data, but L recently, even morphological data can by used P(data | tree)P(tree) P(tree | data) = P(data)

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? • Erected to recognize diversity in Archae,   Ranks  Domain Eubacteria, Eukarya • Major divisions within Eukarya (Plantae,   What do they mean?  Kingdom Animalia, Fungi, etc.)  Are they real?   Phylum • Distinct body plans within Animalia (Plantae  Are they comparable? and Fungi use Divisions)   Class  How should they be used?   Order  Hierarchical framework of • Some sub-groupings  classification  Family  Genus  Genus   Species • ______ Subspecies  Subspecies • Some distinct group within a species?

What are higher taxa and ranks? What are higher taxa and ranks?

 Are they real?  Are they comparable?  Families of Urodela (Salamanders): • 3 domains are monophyletic, no evidence  Domain  Domain that a Domain is something biologically real • Most are monophyletic, no evidence of  Kingdom  Kingdom biological reality  Phylum • Recent evidence suggests that evolution of  Phylum developmental genetic “kernels” defined  Class  Class ______before the Cambrian explosion*  Order  Order  Family • ______of biological reality  Family  Genus  Genus  Species  Species • Most scientists agree that species are actual  Subspecies  Subspecies ______• Little evidence for biological reality Fig. 3-1: Pough et al. 2004. * Davidson & Erwin. 2006. Science 311:796.

What are higher taxa and ranks? An Example – Evolution & Homoplasy in Anolis

• Approximately 400 species of Anolis, ~125 of which  Domain  How should they be used? occur in the Caribbean  Most don’t have a specific meaning,  Kingdom are not biological entities, and aren’t  Phylum comparable to other groups of the same rank  Class  Trend to not use ranks  Order  Including this course!  Family  Use monophyletic groups () to  Genus refer to higher taxa  Species  ______are obviously  Subspecies 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 © J. Losos • The same ______, with similar morphology Crown giant and , have evolved on each of the four big Trunk-crown islands Trunk-crown dwarf © Auburn Herp Soc Cuba

Haiti/Dominican Rep. Trunk-ground

Jamaica Puerto Rico Powell and Russell, 1992. © Reptilien-Center.de

Homoplasy in Anolis – different phylogenetic signal from different datasets Replicate Evolution in Anolis

Locomotor Morphology

mtDNA Anolis ecomorphs evolved in a ______Sequences ______on each of the Greater Antilles

From Jackman et al., 2000. From Jackman et al., 2000.

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