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Tom Horvath Department of Biology SUNY College at Oneonta Not Necessarily on Purpose: Domestication and Speciation in the Canidae Family Tom Horvath Department of Biology SUNY College at Oneonta

Who is Who’s Best Friend? The dog is said to be a man’s (human’s) best friend, but it could also be said that we are a dog’s best friend. Objectives: How to interpret phylogenetic diagrams. Understand how natural selection can work on a population to cause speciation. Distinguish between natural and artificial selection.

Dog Breeds The winner of Best of Show at the 2009 Westminster Dog Show was a Sussex Spaniel. “Stump” the spaniel competed against 170 other breeds of domestic dog (an intraspecies competition). Where do so many breeds come from? How did the species we recognize as the domestic dog arise? Image credit-- Description: Stump is named Best in Show at the Westminster Kennel Club Dog Show. Source: http://www.flickr.com/photos/kjunstorm/3346671755/; also available at http://commons.wikimedia.org/wiki/File:StumpWestminsterShow-wiki.jpg Modifications: Cropped. Author: Kjunstorm (Lori) Licensing: This image is licensed under the Creative Commons Attribution ShareAlike 3.0 License.

The start of our story… A young boy is sitting near the edge of a cave 20,000 years ago. He has just taken out the garbage from the group’s previous day’s activities. The garbage consists of mostly bones and scraps of food from a recent successful hunt. As dusk approaches, the wolves start to arrive. The boy is not frightened. He has seen the wolves many times before. In fact, their arrival is almost ritualistic. They move in from the forest and wait until all the humans have gone into the cave for the night. The boy notices that the same wolf is the first one to get to the good scraps. It doesn’t immediately run off when it sees the boy. Domestic dogs wouldn’t appear on the scene for another 5,000 years….

a boy looked out of a cave … 20,000 years ago, a boy looked out of a cave … Image credits— Description: Cave, National Park Ojców, Poland. Source: http://commons.wikimedia.org/wiki/File:Jaskinia_Ciemna.jpg Author: Jan Jerszyński (2005) Licensing: This image is licensed under the Creative Commons Attribution ShareAlike 2.5 License. Description: Wolf pack (Canis lupus) in animal park of Gramat in France. Source: http://commons.wikimedia.org/wiki/File:Canis_lupus_meute_Gramat.jpg Author: Julien 31 Licensing: This image is licensed under the Creative Commons Attribution ShareAlike 3.0 License.

The Family Canidae Image credit— Source: Anatomical engraving from Handbuch der Anatomie der Tiere für Künstler, Wilhelm Ellenberger and Hermann Baum, and medical illustrator, Hermann Dittrich. 1898 and 1911 through 1925. Scan from http://commons.wikimedia.org/wiki/File:Dog_anatomy_anterior_view.jpg. See also University of Wisconsin Digital Collections at http://digital.library.wisc.edu/1711.dl/Science.VetAnatImgs Licensing: This is a faithful photographic reproduction of an original two-dimensional work of art. The work of art itself is in the public domain its copyright has expired. This applies to the United States, Australia, the European Union and those countries with a copyright term of life of the author plus 70 years.

Members of the Family Canidae Jackal (Black-backed jackal) Fox (Kit fox, Red fox) Wolf (Gray wolf) African wild dog Members of the Family Canidae Image credits— Slide 7 —Top, and Slide 16 —Bottom right Description: Black-backed Jackal in the Masaai Mara, Kenya, April 2008 Source: http://commons.wikimedia.org/wiki/File:Black_Backed_Jackal_Masaai_Mara_April_2008.JPG Author: Simyre Licensing: This image has been released into the public domain by its author Slide 7 —Row 2, left Description: San Joaquin Kit Fox Source: U.S. Fish and Wildlife Service, National Digital Library, http://www.fws.gov/digitalmedia/FullRes/natdiglib/94886A1A-7C7D-4BCD-89E5E6D3F4A65E59.jpg Author: B. Peterson Licensing: Public domain. Slide 7 —Row 2 right, and Slide 16 —Top right Description: Red Fox Source: U.S. Fish and Wildlife Service, National Digital Library, http://www.fws.gov/digitalmedia/FullRes/natdiglib/9C2E1339-AD84-AA2B-DE219D7261B32A0B.jpg Author: Ronald Laubenstein Slide 7 —Row 3, and Slide 16 —Bottom left Description: Gray Wolf, taken in the Garden for Zoologic Research, Tel Aviv University, Israel Source: http://commons.wikimedia.org/wiki/File:TAzooAnimal11.jpg Author: Yonidebest, Hebrew Wikipedia Licensing: The copyright holder of this file, Yonidebest, allows anyone to use it for any purpose, provided that the copyright holder is properly attributed. Slide 7 —Bottom Description: African Wild Dog (Lycaon pictus) Source: http://commons.wikimedia.org/wiki/File:Lycaon_pictus_pg.jpg Author: Philip Gabrielsen Licensing: This image is licensed under the Creative Commons Attribution 2.5 License.

Domestic dogs look like they are more closely related to wolves than other canids. Gray wolf Siberian husky (domestic dog) Image credits— Slide 8 —Left, and Slide 9 —Left Description: Gray wolf, Canis lupus Source: U.S. Fish and Wildlife Service, National Digital Library, http://www.fws.gov/digitalmedia/FullRes/natdiglib/1521AE13-6256-4121-AD32DC27E395E923.jpg Author: Gary Kramer Licensing: Public domain. Slide 8 —Right, Slide 16 —Top left, Description: Siberian Husky Source: http://commons.wikimedia.org/wiki/File:Husky02.jpg Author: Hinrich Licensing: This image is licensed under Creative Commons Attribution ShareAlike 2.0 Germany License. Slide 8 —Lower center Description: Coyote Source: http://commons.wikimedia.org/wiki/File:Coyote2008.jpg Author: Macmanes Licensing: This image is licensed under the Creative Commons Attribution ShareAlike 3.0 License. Coyote

Of course, looks can be deceiving! Gray wolf Domestic dog (Lhasa apso) Image credits-- Slide 9 —Right Description: Lhasa Apso femea com 6 meses Source: http://commons.wikimedia.org/wiki/File:IMG_0258_-_Lhasa_Apso_f%C3%AAmea_de_6_meses.jpg Author: Lcfrederico Licensing: Permission is granted to copy, distribute and/or modify this image under the terms of the GNU Free Documentation License, Version 1.2 or any later version published by the Free Software Foundation; with no Invariant Sections, no Front-Cover Texts, and no Back-Cover Texts. Phylogenetic analyses are more convincing.

Phylogenetic Analysis Phylogenies can be based on morphology Similarity of many morphological characteristics are used (color, size, structure, etc.) Most recent phylogenies are based on molecular similarities E.g., similarities of mitochondrial DNA (mtDNA) sequences More similarities (i.e., fewer dissimilarities) = a closer relationship

The study of evolutionary relationships Phylogeny The study of evolutionary relationships Think of it as a family tree A B C D Recent time Past Remember that this is just a hypothesis of common ancestry. This is rooted. Species A, B, C & D all exist in modern times

The study of evolutionary relationships Phylogeny The study of evolutionary relationships Think of it as a family tree Sp A Sp B Sp C Sp D Unique ancestor of D only Common ancestor=nodes. Blue is a lineage. Common ancestor of both C & D, but not A & B Common ancestor of A, B, C & D

CQ#1: Which statement can be made about this phylogeny? Species A, B, and C are extinct. Species C & D shared a common ancestor more recently than B & D. Species D will display the most advanced morphological characteristics. Species D is most closely related to Species A. Species D evolved from Species C. Sp A Sp B Sp C Sp D

Cladograms are another way to look at phylogenies Image credit— Slide 14 and Slide 29 —Left Description: Domestic dog phylogeny Source: Figure 1 from: Jordana J., Manteca X., Ribo O. Comparative analysis of morphological and behavioral characters in the domestic dog and their importance in the reconstruction of phylogenetic relationships in canids. Genet. Mol. Biol. [serial on the Internet]. 1999 Mar [cited 2009 July 24] ; 22(1): 49–57. Available from: http://www.scielo.br/scielo.php?script=sci_arttext&pid=S1415-47571999000100011&lng=en. doi: 10.1590/S1415-47571999000100011. Permission: According to http://www.scielo.br/revistas/gmb/iaboutj.htm, articles may be reproduced without specific permission from Genet. Mol. Biol. for use on a non-profit basis in teaching.

Species A Species B Species C Species D Species E Species F Introduce clades here. Clades share a derived characteristic. Also called synapomorphies. Species A and B are sister taxa Species A & B are more similar to each other than they are to any other species Species A-D are more similar to each other than they are to either species E or F Species C & D shared a common ancestor in more recent times than the shared common ancestor of A-D

In Groups: Discuss relationships and create a cladogram for: Fox Domestic dog Image credit-- Wolf Jackal

CQ#2: Which of the following most closely resembles your cladogram? Wolf Jackal Fox Dog Wolf Jackal Fox A. C. D. Dog Wolf Jackal Fox Dog Wolf Jackal Fox E. Wow, mine looks like none of these

Canid Phylogeny Slide 18 —Center, Slide 19 —Right Description: Canid phylogeny Source: http://whozoo.org/mammals/Carnivores/canid_phylogeny.jpg Permission: According to http://whozoo.org/copyright.htm, “Images may be downloaded and reproduced for non-profit personal or educational purposes without further permission.”

CQ#3: According to the molecular evidence shown in the cladogram, which statement is most true? Foxes and wolves are closely related. Domestic dogs and wolves are as closely related to each other as they are to coyotes. Out of the canids tested, a Gray wolf is the most likely ancestor of the domestic dog. The domestic dog is the most evolved of the canids.

Comparing wolves with dogs Morphological comparisons (examples) Dogs tend to have curled tails, wolves have straight tails. Dogs tend to have smooth short coats. Skull shape differs. Molecular comparisons Gray wolves and dogs differ by no more than 0.2% in their mtDNA sequence. In contrast, gray wolves and coyotes differ by at least 4%.

How did dogs evolve from wolves? Competing hypotheses Ancestral wolf pups were domesticated intentionally by early humans – Artificial selection. Ancestral wolf populations experienced natural selection forces that favored dog-like characteristics.

Hypotheses 1 – Artificial Selection Arguments for It makes intuitive sense that ancestral wolves could be domesticated easily since they are so dog-like. Early humans would have intentionally bred ancestral wolves. Arguments against Why would any human want to deal with an animal that avoids humans? Modern wolves can not be “domesticated” by training alone. It takes intensive and sophisticated selective breeding.

Hypothesis 2 – Natural Selection Canids are very resourceful & would have found human waste piles good foraging – wolves were living close to humans. Wolves are shy, skittish animals – only “adventurous” wolves would have stayed close to the waste piles while humans were around. The “adventurous” wolves were the best fed and hence had high fitness. This hypothesis is based on Coppinger, Raymond and Lorna Coppinger Dogs: A New Understanding of Canine Origin, Behavior and Evolution. ISBN: 978-0-226-11563-4 (ISBN-10: 0-226-11563-1) Fall 2002

CQ#4: What is fitness in the evolutionary sense? Being the strongest and most able to get food. Being able to survive and reproduce more successfully than others. Being the most aggressive and fending off potential predators. Being the smartest and remembering where the best food resources are.

CQ#5: What characteristic is being selected for in the ancestral wolf population under Hypothesis 2? Ability to interact with humans. Capability of eating human waste. Behaviors that lead to tolerance of humans being around. Inability to run away from humans.

A Plausible Series of Events Those ancestral wolves that tended to tolerate humans would have had the best access to high-quality food (scraps). Canids likely would have protected their scrap piles from other invading animals and possibly even strange humans.

What did humans give dogs? Easy access to high-quality food. A safe “home” to raise pups. What else… The bullets could be left out and allow students to group and come up with a list of advantages that humans provided for dogs (or ancestral wolves). Image credit— Slide 27 Description: Two dogs on sofa Source: Thomas Horvath, Department of Biology, SUNY College at Oneonta Permission: Used with permisssion.

CQ#6: What did the first "dogs" give humans that is the most significant in evolutionary terms? An efficient garbage disposal. An affectionate pet. An early warning system that someone or something was approaching. This could be an optional slide Alternative is to ask students to re-group and come up with evolutionary advantages that humans received by their association with dogs.

Phylogeny of Domestic Dog Breeds – Result of Artificial Selection Image credits— Slide 29 —Top left Description: Afghan Hound Source: http://commons.wikimedia.org/wiki/File:Afghan_Hound.jpg Author: Sannse Permission: Permission is granted to copy, distribute and/or modify this document under the terms of the GNU Free Documentation License, Version 1.2 or any later version published by the Free Software Foundation; with no Invariant Sections, no Front-Cover Texts, and no Back-Cover Texts. Slide 29 —Top right Description: Irish Setter Source: http://commons.wikimedia.org/wiki/File:Can_Setter_dog_GFDL.jpg Author: Luis Miguel Bugallo Sánchez Slide 29 —Farthest left Description: Akita-Inu Source: http://commons.wikimedia.org/wiki/File:Akita_inu.jpeg Author: B@rt Slide 29 —Middle Description: Alaskan Malamute Source: http://he.wikipedia.org/wiki/%D7%A7%D7%95%D7%91%D7%A5:Alaskanmalamute0b.jpg Author: PardoY Slide 29 —Farthest right Description: Siberian Husky Source: http://commons.wikimedia.org/wiki/File:Siberian-husky.jpg Author: Utopialand Slide 29 —Lower left Description: Samoyed Source: http://commons.wikimedia.org/wiki/File:Samoyed_600.jpg Slide 29 —Lower right Description: Chow-chow Source: http://commons.wikimedia.org/wiki/File:Chowchow.jpg Author: Jurriaan Schulman

Evidence for Evolution Paleontology – fossils show change in a species over time Biogeography – Similar species are found in similar ecosystems around the world Morphology – Comparing structures Homologous structures – body parts with similar structure but possible different function. Shows common ancestry Analogous structures – similar structure develops in organisms that share a common ecosystem but not a common ancestry Biochemical or Molecular - Similarities in gene sequences, proteins, DNA

Fossil record Sedimentary rock are richest source of fossils fossil record is a substantial, but incomplete, chronicle of evolutionary history incomplete historical documents of biology history of life on Earth is punctuated by mass extinctions 5000 year old ice mummy found on an Alpine ridge dividing Austria from Italy at 10,500 feet above sea level. 2004-2005

Relative Dating

Evolutionary Time Scale Microevolution – changing of allele frequencies in a population over time. Macroevolution – patterns of change over geologic time. Determines phylogeny Gradualism – species are always slowly evolving Punctuated equilibrium – periods of massive evolution followed by periods with little to no evolution

Homologous Structures Anatomical evidence Anatomical evidence 1. These structures are called… homologous 2. These structures are evidence for…. common ancestry 3. similar internal structure = similar development 4. different function = different environment & niche 5. close evolutionary r’ship!

Analogous structures Solving a similar problem with a similar solution Convergent evolution Don’t be fooled by their looks! same function different development & anatomy Those fins & tails & sleek bodies are analogous structures! convergent evolution - NOT common ancestor Those fins & tails & sleek bodies are analogous structures! Does this mean they have a recent common ancestor? Solving a similar problem with a similar solution

Evaluating molecular homologies Aligning DNA sequences more bases in common = more closely related analyzed by software 2004-2005 beware of molecular homologies

Comparative hemoglobin structure Molecular Homology The sequence in DNA & proteins is a molecular record of evolutionary relationships. Why compare DNA & proteins across species? 10 20 30 40 50 60 70 80 90 100 110 120 Lamprey Frog Bird Dog Macaque Human 32 8 45 67 125 Comparative hemoglobin structure Molecular Record What are we comparing here? - comparing DNA (base sequence) & proteins (amino acid sequence) What assumption do we make about genes and relatedness?  the more closely related, the more DNA bases & amino acids you have in common have to compare genes for protein the organisms have in common… can’t compare genes for proteins you don’t have Number of amino acid differences between hemoglobin (146 aa) of vertebrate species and that of humans

Phylogeny & Systematics evolutionary history of a species based on common ancestries inferred from fossil record morphological & biochemical resemblances molecular evidence Systematics connects classification system to phylogeny by categorizing & naming organisms 2004-2005

Systematics Connecting classification to phylogeny hierarchical system Carolus Linnaeas latin binomial genus species 2004-2005

Illustrating phylogeny Cladograms patterns of shared characteristics Classify organisms according to the order in time at which branches arise along a phylogenetic tree 2004-2005

Molecular Systematics Hypothesizing phylogenies using molecular data apply principle of parsimony simplest explanation fewest evolutionary events that explain data hypothetical bird species 3 possible phylogenies (there are more) 2004-2005

Parsimony Choose the “tree” that explains the data invoking the fewest number of evolutionary events

Parsimony & analogy vs. homology Phylogenetic trees are hypotheses Which is the most parsimonious tree? 2004-2005

Modern Systematics Shaking up some trees! Crocodiles are now thought to be closer to birds than other reptiles 2004-2005

TAXA Lancelet (outgroup) (outgroup) Lancelet Lamprey Leopard Bass Frog Figure 20.11 TAXA Lancelet (outgroup) (outgroup) Lancelet Lamprey Leopard Bass Frog Turtle Lamprey Vertebral column (backbone) 1 1 1 1 1 Bass Vertebral column Hinged jaws 1 1 1 1 Frog Four walking legs Hinged jaws CHARACTERS 1 1 1 Turtle Four walking legs Amnion 1 1 Figure 20.11 Constructing a phylogenetic tree Amnion Hair 1 Leopard Hair (a) Character table (b) Phylogenetic tree 46

Three Big Ideas about Phylogenetic Trees Time runs from the root to the tips of the tree, not across its tips The branching pattern of a tree indicates relatedness—taxa that share more recent common ancestors are more closely related. Trees depict evolutionary relationships, not evolutionary progress. Try #8 on page 398.