67
1 Systematics, Phylogenies, and Comparative Biology Chapter 23

Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

  • Upload
    others

  • View
    2

  • Download
    0

Embed Size (px)

Citation preview

Page 1: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

1

Systematics,

Phylogenies,

and Comparative

Biology

Chapter 23

Page 2: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Systematics

• All organisms share many

characteristics:

– Composed of one or more

cells

– Carry out metabolism

– Transfer energy with ATP

– Encode hereditary

information in DNA

• Tremendous diversity of life

– Bacteria, whales, sequoia

trees 2

http://www.pbs.org/wgbh/nova/evolution/ten-great-advances-evolution.html

Page 3: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

3

• Biologists group organisms based on shared

characteristics and newer molecular sequence

data

– Since fossil records are not complete,…

– Scientists rely on other types of evidence to establish

the best hypothesis of evolutionary relationships

Actual

fossil

record

Fairly

complete

record

Poor

fossil

record

Misinterpreted

fossil record

http://www.mun.ca/biology/scarr/2900_Incomplete_Fossil_Record.htm

Page 4: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

4

• Systematics

– Reconstruction and study

of evolutionary

relationships

• Phylogeny

– Hypothesis about patterns

of relationship among

species

– Cladogram or

phylogenetic tree

http://paleocave.sciencesortof.com/2010/12/diagraming-evolution-or-how-to-read-a-cladogram/

Page 5: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

5

Darwin envisioned that all

species were descended

from a single common

ancestor

• He depicted this history of

life as a branching tree

• “Descent with modification”

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

a.

Reproduced by kind permission of the Syndics of Cambridge University Library, Darwin’s Notebook ‘B’,

‘Tree of Life’ Sketch, p. 36 from DAR.121 D312

Page 6: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

6

• Key to interpreting a phylogeny – look at how

recently species share a common ancestor

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Variations of a Cladogram

Gibbon Human Chimp Gorilla Orangutan Gibbon Orangutan Gorilla Human Chimp

Chimp

Human

Gorilla

Orangutan

Gibbon

1

3

1

2

2

3

3

2

1

Version 1 Version 2

b.

Version 3

Page 7: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Similarity may not accurately predict

evolutionary relationships

– Early systematists relied on the

expectation that the greater the time

since two species diverged from a

common ancestor, the more different

they would be

– Rates of evolution vary

• Evolution may not be unidirectional

– Evolution is not always divergent

• Convergent evolution – similar characters

not derived from common ancestery 7

http://matthewbonnan.wordpress.com/2012/06/02/lets-face-it-birds-are-dinosaurs-part-1/

Page 8: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Cladistics

• Derived characteristic – Similarity that is inherited from the

most recent common ancestor of an entire group

• Ancestral characteristic – Similarity that arose prior to the

common ancestor of the group

• In cladistics, only shared derived characters (synapomorphies) are considered informative about evolutionary relationships

8 http://www.nature.com/scitable/topicpage/trait-evolution-on-a-

phylogenetic-tree-relatedness-41936

Page 9: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Characters can be any aspect of the phenotype – Morphology – Physiology

– Behavior – DNA; RNA

• Characters should exist in recognizable character states – Presence vs. absence

– Example: Character “teeth” in amniote vertebrates has two states, present in most mammals and reptiles, and absent in birds and turtles

9

http://sciencenotes.wordpress.com/2008/05/10/

Page 10: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Examples of ancestral versus

derived characters, often

dependent on context

– Presence of hair is a shared

derived feature of mammals

– Presence of lungs in mammals is

an ancestral feature; also present

in amphibians and reptiles

– Shared, derived feature of hair

suggests that all mammal

species share a common

ancestor that existed more

recently than the common

ancestor of mammals,

amphibians, and reptiles

10

http://sciencenotes.wordpress.com/2008/05/10/

Page 11: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Manual cladistic analysis

• First step is to polarize the characters

(are they ancestral or derived)

– Example: polarize “hair” means to

determine presence or absence in the

most recent common ancestor

– Outgroup comparison used

• Species or group of species that is closely

related to, but not a member of, the group

under study is designated as the outgroup

• Outgroup species do not always exhibit the

ancestral condition 11

Page 12: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

12

“1” = possession of derived character state

“0” = possession of ancestral character state

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Traits:

Organism

Jaws Lungs Amniotic

Membrane Hair No Tail Bipedal

Lamprey

Shark

Salamander

Lizard

Tiger

Gorilla

Human

0 0 0 0 0 0

1

1

1

1

1

1

0

1

1

1

1

1

0

0

1

1

1

1

1 1 1

1 1 0

1 0 0

0 0 0

0

0 0 0

0 0

a.

Page 13: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

13

• Derived characters between the cladogram branch points are

shared by all organisms above the branch points and are not

present in any below them

• The outgroup (in this case, the lamprey) does not possess any

of the derived characters

Outgroup may or may not exhibit ancestral form

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Lamprey Shark Salamander Lizard Tiger Gorilla Human

Jaws

Lungs

Amniotic

membrane

Hair

Tail loss

Bipedal

b.

Page 14: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• When the group under study exhibits multiple

character states, and one of those states is exhibited

by the outgroup, then that state is ancestral and

other states are derived

– Most reliable if character state is exhibited by several

different outgroups

– Presence of teeth in mammals and reptiles is ancestral

– Absence of teeth in birds and turtles is derived

14

http://ncse.com/book/export/html/11798

Page 15: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Cladogram

– Depicts a hypothesis of evolutionary relationships

• Clade

– Species that share a common ancestor as indicated

by the possession of shared derived characters

– Evolutionary units refer to a common ancestor and

all descendants

– Synapomorphy – derived character shared by

clade members

15

http://www.vanderbilt.edu/peabody/novick/evol_diagrams.html

Page 16: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Simple cladogram is a nested set of clades,

each characterized by its own synapomorphies

– Amniotes are a clade for which the evolution of an

amniotic membrane is a synapomorphy

– Within that clade, mammals are a clade, with hair as

a synapomorphy

16

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Lamprey Shark Salamander Lizard Tiger Gorilla Human

Jaws

Lungs

Amniotic

membrane

Hair

Tail loss

Bipedal

b.

Page 17: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Plesiomorphies – ancestral

states

• Symplesiomorphies –

shared ancestral states

• Character state “presence

of a tail” – Exhibited by lampreys, sharks,

salamanders, lizards, and

tigers

– Are tigers more closely related

to lizards and sharks than

apes and humans?

– Symplesiomorphies reflect

character states inherited from

a distant ancestor, they do not

imply that species exhibiting

that state are closely related 17

Salamander Frog Lizard Tiger Gorilla Human

Tail loss

Amniotic

membrane

Hair

Tail loss

a.

Page 18: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Homoplasy – a shared character state that

has not been inherited from a common

ancestor

– Convergent evolution (independent evolution)

– Evolutionary reversal (return to ancestral state)

18

Modified from… http://palaeoblog.blogspot.com/2011_02_01_archive.html

Homoplasy –

Convergence

A derived trait

present in 2

lineages that lack

common ancestor

Homoplasy – Reversal

Trait in ancestor but not

its immediate

descendents, but

appears later in

subsequent descentdent

Homology

Trait present in an

ancestor and in

itsdescentdents

Page 19: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Systematists rely on the principle of parsimony,

which favors the hypothesis that requires the

fewest assumptions

19

http://undsci.berkeley.edu/article/howscienceworks_12

Page 20: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

20

Based on the principle of parsimony, the cladogram that

requires the fewest number of evolutionary changes is

favored

• The cladogram in (a) requires four changes,

whereas…

• (b) requires five

Salamander Frog Lizard Tiger Gorilla Human Salamander Lizard Tiger Frog Gorilla Human

Tail loss

Amniotic

membrane

Hair

Tail loss

Amniotic

membrane

Hair

Tail loss

Amniotic

membrane

loss

Hair loss

a. b.

Page 21: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Systematists increasingly

use DNA sequence data

to construct phylogenies

• Large number of

characters that can be

obtained through

sequencing

• Character states are

polarized by reference to

the sequence of an

outgroup

• Cladogram is constructed

that minimizes the amount

of character evolution

required

21

http://www.biocompare.com/Application-Notes/41940-Mutation-Detection-For-The-K-Rasand-P16-Genes/

Page 22: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

22

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

G

Site

Species A

Species B

Species C

Species D

Outgroup

DNA Sequence

1 2 3 4 5 6 7 8 9 10

A T A T C C G T A T

A C A G C C G C A T

A C A T C G G T G G

G C A T A G T G T

G C A T A G G C G T

Page 23: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

23

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Outgroup Species B Species D Species A Species C

Homologous

evolutionary

changes

Homoplastic

evolutionary

changes

2:T C

4:T

8:T

G

C

10:T G

8:T C

9:A

6:C

G

G

5:C A

1:A G

Parsimony also attempts to minimize homoplasies

Page 24: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Other Phylogenetic Methods

Potential problems with DNA sequences… • Some characters evolve rapidly, so using

parsimony principle may be misleading

• Stretches of DNA with no function have high rates of evolution of new character states due to genetic drift

• Only 4 character states are possible (A, T, G, C)

so there is a high probability that two species will

independently evolve the same derived character

state at any particular base position… appear

homologous when they are not

24

Page 25: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Statistical (maximum likelihood) approach

– Start with an assumption about the rate at which

characters evolve (change)

– Fit the data to these models to derive the phylogeny

that best follows (i.e., “maximally likely”) these

assumptions

– Narrows down possibile cladograms to “most likely”

– Usually compared with parsimony model

25

Other Phylogenetic Methods

http://www.scielo.br/scielo.php?pid=s1415-47571999000300009&script=sci_arttext

Maximum parsimony tree Maximum likelihood tree

Page 26: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Molecular clock

– Assumes rate of evolution of a molecule is constant

through time (see figure)

– Divergence in DNA can be used to calculate the

times at which branching events have occurred

– Mutation rates may not be constant, depending on

selection pressures on traits

26

Other Phylogenetic Methods

http://www.mun.ca/biology/scarr/Molecular_Clock.html

Page 27: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Systematics and Classification • Classification

– How we place species and higher groups into the

taxonomic hierarchy – should follow evolutionary

history

– Genus, family, order, class, phylum, kingdom, domain

• Monophyletic group (the “goal”)

– Includes the most recent common ancestor of the

group and all of its descendants (clade)

27

a.

Archosaurs

Monophyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

Page 28: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Systematics and Classification • Monophyletic group

– Includes the most recent common ancestor of the

group and all of its descendants (clade)

– “Single cut” of cladogram

– The goal of “good” phylogeny

28 a.

Archosaurs

Monophyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

Page 29: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

29

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

a.

Archosaurs

Monophyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

Page 30: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Systematics and Classification

• Paraphyletic group

– Includes the most

recent common

ancestor of the group,

but not all its

descendants

• Polyphyletic group

– Does not include the

most recent common

ancestor of all

members of the group

30

b.

Dinosaurs

Paraphyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

c.

Polyphyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

Flying Vertebrates

Page 31: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Taxonomic hierarchies

are based on shared

traits, should reflect

evolutionary relation-

ships, i.e. be

monophyletic

• Birds as an example

– Must be included in

dinosaurs to be

monophyletic

31

b.

Dinosaurs

Paraphyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

a.

Archosaurs

Monophyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

Page 32: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

32

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

b.

Dinosaurs

Paraphyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

Page 33: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

33

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

c.

Polyphyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

Flying Vertebrates

Page 34: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

34

• The traditional classification included two groups that we

now realize are not monophyletic: the green algae and

bryophytes

– New classification of plants does not include these groups

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Red Algae Green Algae

Chlorophytes Charophytes Liverworts

Bryophytes

Hornworts Mosses

Vascular Plants

Page 35: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Biological species concept (BSC) – Defines species as groups of

interbreeding populations that are reproductively isolated

• Phylogenetic species concept (PSC)

– Species is a population or set

of populations characterized

by one or more shared derived

characters

35

Species concepts

http://evolution.berkeley.edu/evosite/evo101/VA1BioSpeciesConcept.shtml

Page 36: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

36

• PSC solves 2 BSC

problems

– BSC cannot be applied to

allopatric (separate)

populations – would they

interbreed?

• PSC looks to the past to see if

they have been separated

long enough to develop their

own derived characters

– BSC can be applied

only to sexual species • PSC can be applied to both

sexual and asexual species

http://evolution.berkeley.edu/evosite/evo101/VA1BioSpeciesConcept.shtml

Page 37: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• PSC still controversial

– Critics contend it will lead to the recognition of

even slightly different populations as distinct

species – “splitting”

– Paraphyly problem – if species C evolved

apart from other genetically connected

“species” A,B, D & E

37

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

A B C D E

Page 38: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Phylogenetics

• Basis for all comparative biology

• Homologous structures

– Derived from the same ancestral source

– Dolphin flipper and horse leg

38

http://bytesizebio.net/2009/07/15/distant-homology-and-being-a-little-pregnant/

Page 39: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Phylogenetics

• Homoplastic structures are not

– Analogous – similar function due to similar selection pressures

– Wings of birds and dragonflies

• Phylogenetic analysis can help determine which structures are homologous vs. analogous

39

http://www.bio.miami.edu/dana/106/106F05_4.html

This is a moth

Page 40: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Parental care in dinosaurs

initially treated as unexpected

– Examination of phylogenetic

comparison of dinosaurs

indicates they are most closely

related to crocodiles and birds –

both show parental care

– Parental care in three groups not

convergent (or analogous) but

homologous behaviors Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

a. a: Image #5789, photo by D. Finnin/American Museum of Natural History

b. 40 b: © Roger De La Harpe/Animals Animals

Fossil dinosaur

incubating eggs

a.

Archosaurs

Monophyletic Group

Bat Crocodile Stegosaurus Hawk Giraffe Turtle Tyrannosaurus Velociraptor

Page 41: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Homoplastic convergence

• Ex: saber teeth

– Evolved independently in different clades of

extinct carnivores

– Similar body proportions (cat) & predatory lifestyle

– Most likely evolved independently at least 3 times

41

Monotremes

Marsupials Placentals

Phylogeny of Mammals

Carn

ivo

res

Sab

er-

too

thed

mars

up

ial

Nimravids are a now-extinct group of catlike carnivores

Carnivores

Felines

Nimravids

Sab

er-

too

thed

cat

Hyen

as

Civ

ets

Mo

ng

oo

ses

Sab

er-

too

thed

nim

rav

id

Phylogeny of Carnivores

Bears

, seals

,

weasels

, can

ids,

an

d r

acco

on

s

Sab

er-

too

thed

nim

rav

id

Monotremes

Marsupials Placentals

Phylogeny of Mammals

Carn

ivo

res

Sab

er-

too

thed

mars

up

ial

Page 42: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

42

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Monotremes

Marsupials Placentals

Phylogeny of Mammals

Carn

ivo

res

Sab

er-

too

thed

ma

rsu

pia

l

Page 43: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

43

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Nimravids are a now-extinct group of catlike carnivores

Carnivores

Felines

Nimravids

Sa

be

r-to

oth

ed

cat

Hye

na

s

Civ

ets

Mo

ng

oo

ses

Sa

be

r-to

oth

ed

nim

ravid

Phylogeny of Carnivores

Be

ars

, s

ea

ls,

wea

se

ls, c

an

ids

,

an

d r

ac

co

on

s

Sa

be

r-to

oth

ed

nim

ravid

Page 44: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Homoplastic convergence

• Ex: plant conducting tubes

– Sieve tubes facilitate long-

distance transport of food and

other substances in vascular

plants

• Essential to the survival of

tall plants on land

– Brown algae also have sieve

elements

– Closest ancestor of these two

taxa is a single-celled organism

that could not have had a

multicellular transport system… 44

http://www.cartage.org.lb/en/themes/sciences/botanicalsciences/plantsstructure/plantsstructure/plantsstructure.htm

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Page 45: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

45

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

(left): © Lee W. Wilcox; (right): © Dr. Richard Kessel & Dr. Gene Shih/Visuals Unlimited

– Closest ancestor of these two taxa is a single-celled

organism that could not have had a multicellular

transport system

µ m 2 µ m

Stramenopiles Alveolates Red Algae Chlorophytes Chlorophytes Liverworts Hornworts Mosses Tracheophytes

Brown Alga

60

Angiosperms

Page 46: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Comparative Biology

Most complex characters do not evolve in one step

• Evolve through a sequence of evolutionary changes

• Initial stages of a character evolved as an adaptation to some environmental selective pressure

• Modern-day birds are exquisite flying machines – Wings, feathers, light bones,

breastbone 46

http://en.wikipedia.org/wiki/White_feather

Page 47: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• First featherlike structure evolved in theropod phylogeny – For insulation or perhaps decoration

– Selection works with what is present 47

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Other dinosaurs Coelophysis Tyrannosaurus Sinosauropteryx Velociraptor Caudipteryx Archaeopteryx Modern Birds

Light bones

Wishbone, breastbone,

loss of fingers 4 and 5

Downy feathers

Long arms, highly mobile wrist, feathers

with vanes, shafts, and barbs

Long, aerodynamic

feathers

Arms longer than legs

Loss of teeth and

reduction of tail

http://news.nationalgeographic.com/news/2011/09/pictures/110915-

amber-dinosaur-feathers-color-science-birds-alberta/

70-85 my old

feathers trapped

in amber

Page 48: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Comparative Biology

• Ex: Vertebrate eye vs.

cephalopod eye

– Both “camera” eyes,

look similar

– Developmental process

is very different

– Highly complex

structures,

independently derived

(convergence)

48

http://www.pnas.org/content/94/6/2098/F1..

Page 49: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Phylogenetic methods can be

used to distinguish between

competing hypotheses

• Larval dispersal in marine

snails

– Some snails produce

microscopic larvae that drift

in the ocean currents

– Some species have larvae

that settle to the ocean

bottom and do not disperse

49

http://www.pbs.org/wgbh/nova/sharks/island/underparksveliger.html

Veliger larvae

dispersal stage

Jump to

Slide #59

Phylogenetic Analysis

Page 50: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Fossil record shows an increase in non-

dispersing snails

50

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Eocene

Percentage of species with

nondispersing larvae

40 MYA

45 MYA

50 MYA

55 MYA

60 MYA

65 MYA

0 100 50

Page 51: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

51

• Two processes could produce an increase in non-dispersing larvae – Evolutionary change from

dispersing to non-dispersing occurs more often than change in the opposite direction

– Species that are non-dispersing speciate more frequently, and/or become extinct less frequently than dispersing species

• The two processes would result in different phylogenetic patterns

a.

Dispersing larvae

Nondispersing larvae

b.

Dispersing larvae

Nondispersing larvae

Page 52: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• In this hypothetical example, the evolutionary

transition from dispersing to nondispersing larvae

occurs more frequently (four times) than the converse

(once)

52

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

a.

Dispersing larvae

Nondispersing larvae

Page 53: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• By contrast, clades that have nondispersing larvae

diversify to a greater extent due to higher rates of

speciation or lower rates of extinction (assuming that

extinct forms are not shown)

53

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

b.

Dispersing larvae

Nondispersing larvae

Page 54: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Phylogeny for Conus, a genus of marine snails.

– Nondispersing larvae have evolved eight separate

times from dispersing larvae, with no instances of

evolution in the reverse direction.

– This phylogeny does not show all species, however;

nondispersing clades contain on average 3.5 times as

many species as dispersing clades.

54

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

c.

Dispersing larvae Nondispersing larvae

Dispersing larvae

Nondispersing larvae

Page 55: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Analysis indicates:

– Evolutionary increase in non-dispersing larvae

through time may be a result of both as a bias in the

evolutionary direction and an increase in rate of

diversification (more likely to be isolated higher

speciation rate)

– Lack of evolutionary reversals not surprising…

evolution of non-dispersing larvae a one-way street

55

c.

Dispersing larvae Nondispersing larvae

Page 56: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Loss of larval stage in marine

invertebrates

– Eggs develop directly into

adults

– Nonreversible evolutionary

change

– Marine limpets: show direct

development has evolved

many times

– Three cases where evolution

reversed and larval stage re-

evolved

– Evidence in second species

supports hypotheses 56

a.

Larval stage

Direct development

Page 57: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

57

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

a.

Larval stage

Direct development

Page 58: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Less parsimonious

interpretation of

evolution in the clade in

the light blue box is

that, rather than two

evolutionary reversals,

six instances of the

evolution of

development occurred

without any

evolutionary reversal

58

b.

Larval stage

Direct development

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Page 59: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Phylogenetics helps explain species diversification

• Use phylogenetic analysis to suggest and test hypotheses

• Insight into beetle diversification – Correspondence between phylogenetic position and

timing of plant origins suggests beetles are remarkably conservative in their diet

59

Beetle box: 102 species of

beetles from Madagascar,

representing ~0.025% of all

described beetle species

http://debivort.org/blog/2010/08/01/beetle-box/

Phylogenetic Analysis

Page 60: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

60

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Tertiary

Cretaceous

Jurassic

Triassic Angiosperm

Cycad

Conifer

Number

of Species 85

2000

24

1500

41

,60

2

10

20

750

78

25,0

00

3

400

8

1 8

33

,40

0

– Beetle families that specialize on conifers have the deepest branches

– Beetles that specialize on Angiosperms have shorter branches

Page 61: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• Phylogenetic explanations for beetle diversification – It is not the evolution of

herbivory itself

– Specialization on angiosperms was prerequisite for diversification

– Specialization has arisen five times independently within herbivorous beetles

– Angiosperm-specializing clade is more species-rich than the clade most closely related

61

http://meetthespeciesdotorg.wordpress.com/2012/08/23/sarah-beynon-on-beetles/

Tertiary

Cretaceous

Jurassic

Triassic Angiosperm Cycad Conifer

Number

of Species

85

2000

24

1500

41,6

02

10

20

750

78

25,0

0

0 3

40

0

8

1 8

33,4

00

Page 62: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Disease Evolution

AIDS first recognized in 1980s

• Current estimate: > 33 million people infected with human immunodeficiency virus (HIV); > 2 million die each year

• Simian immunodeficiency virus (SIV) found in 36 species of primates – Does not usually cause illness in

monkeys

– Around for more than a million years as SIV in primates 62

http://www.sciencenews.org/view/access/id/46079/description/Back_story_Untangling_SIV_and_HIV-1

Page 63: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

Phylogenetic analysis of HIV and SIV

1. HIV descended from SIV

– All strains of HIV are nested within clades of SIV

2. Several different strains of HIV exist – Independent transfers from

different primate species

– Each human strain is more closely related to a strain of SIV than to other HIV strains

• Separate origins for HIV strains 63

HIV

- 1 g

rou

p M

HIV

- 1 g

rou

p N

SIV

- ch

imp

an

zee

HIV

- 1 g

rou

p O

SIV

- d

rill

SIV

- ta

nta

lus

SIV

- so

oty

man

gab

ey

HIV

- 2 g

rou

p A

SIV

- ch

imp

an

zee

SIV

- ch

imp

an

zee

SIV

- ch

imp

an

zee

SIV

- ch

imp

an

zee

HIV

- 1 g

rou

p P

SIV

- ch

imp

an

zee

SIV

- re

d-c

ap

ped

man

gab

ey

SIV

- g

ori

lla

SIV

- verv

et

mo

nkey

HIV

- 2 g

rou

p B

SIV

- syke’s

mo

nkey

SIV

- D

e B

razzas m

on

key

SV

- so

oty

man

gab

ey

SV

- g

reate

r sp

ot-

nasal

mo

nkey

Indicates

transmission to humans

Page 64: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

3. Humans have acquired

HIV from different host

species

– HIV-1, which is the virus

responsible for the global

epidemic, has three

subtypes

• Each subtype is most

closely related to a

different strain of

chimpanzee SIV,

indicating that the

transfer occurred from

chimps to humans

64

HIV

- 1

gro

up

M

HIV

- 1

gro

up

N

SIV

- c

him

pa

nze

e

HIV

- 1

gro

up

O

SIV

- d

rill

SIV

- ta

nta

lus

SIV

- s

oo

ty m

an

ga

be

y

HIV

- 2

gro

up

A

SIV

- c

him

pa

nze

e

SIV

- c

him

pa

nze

e

SIV

- c

him

pa

nze

e

SIV

- c

him

pa

nze

e

HIV

- 1

gro

up

P

SIV

- c

him

pa

nze

e

SIV

- re

d-c

ap

pe

d m

an

ga

be

y

SIV

- g

ori

lla

SIV

- ve

rve

t m

on

ke

y

HIV

- 2

gro

up

B

SIV

- s

yk

e’s

mo

nk

ey

SIV

- D

e B

razza

s m

on

ke

y

SV

- s

oo

ty m

an

ga

be

y

SV

- g

rea

ter

sp

ot-

na

sa

l m

on

ke

y

Indicates

transmission to humans

Page 65: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

– Subtypes of HIV-2, which is

much less widespread, are

related to SIV found in West

African monkeys, primarily

the sooty mangabey

(Cercocebus atys)

• Moreover, the subtypes of

HIV-2 also appear to

represent several

independent cross-

species transmissions to

humans

65

HIV

- 1

gro

up

M

HIV

- 1

gro

up

N

SIV

- c

him

pa

nze

e

HIV

- 1

gro

up

O

SIV

- d

rill

SIV

- ta

nta

lus

SIV

- s

oo

ty m

an

ga

be

y

HIV

- 2

gro

up

A

SIV

- c

him

pa

nze

e

SIV

- c

him

pa

nze

e

SIV

- c

him

pa

nze

e

SIV

- c

him

pa

nze

e

HIV

- 1

gro

up

P

SIV

- c

him

pa

nze

e

SIV

- re

d-c

ap

pe

d m

an

ga

be

y

SIV

- g

ori

lla

SIV

- ve

rve

t m

on

ke

y

HIV

- 2

gro

up

B

SIV

- s

yk

e’s

mo

nk

ey

SIV

- D

e B

razza

s m

on

ke

y

SV

- s

oo

ty m

an

ga

be

y

SV

- g

rea

ter

sp

ot-

na

sa

l m

on

ke

y

Indicates

transmission to humans

Page 66: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

66

Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

HIV

- 1

gro

up

M

HIV

- 1

gro

up

N

SIV

- c

him

pa

nzee

HIV

- 1

gro

up

O

SIV

- d

rill

SIV

- ta

nta

lus

SIV

- s

oo

ty m

an

gab

ey

HIV

- 2

gro

up

A

SIV

- c

him

pa

nzee

SIV

- c

him

pa

nzee

SIV

- c

him

pa

nzee

SIV

- c

him

pa

nzee

HIV

- 1

gro

up

P

SIV

- c

him

pa

nzee

SIV

- re

d-c

ap

ped

ma

n g

ab

ey

SIV

- g

ori

lla

SIV

- ve

rve

t m

on

ke

y

HIV

- 2

gro

up

B

SIV

- s

yk

e’s

mo

nke

y

SIV

- D

e B

razza

s m

on

ke

y

SV

- s

oo

ty m

an

gab

ey

SV

- g

rea

ter

sp

ot-

na

sa

l m

on

ke

y

Indicates

transmission to humans

Page 67: Systematics, Phylogenies, and Comparative Biologywtyler-irsc.weebly.com/uploads/1/9/8/6/19866611/chapt_23...• Examples of ancestral versus derived characters, often dependent on

• HIV mutates so rapidly that

a single HIV-infected

individual often contains

multiple genotypes in a

patient’s body

• As a result, it is possible to

create a phylogeny of HIV

strains and to identify the

source of infection of a

particular individual

• In this case, the HIV strains

of the victim (V) clearly are

derived from strains in the

body of another individual,

the patient 67 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Vic

tim

Pati

en

t

Co

mm

un

ity