Elements of appendicular skeleton:

f
forelimb
li b
Pectoral girdle:

F
Forelimb:
m

Mammals
Anterior

Posterior

Limbs
Li b and
d associated
i t d muscles
l are
derived from somatic hypomere:

Hox gene expression in:

Bony fishes

Tetrapods

Pectoral girdle

Modifications of
the basic fish
pattern in
vertebrates

Anterior

Dermal
elements

Endochondral
elements

Shark

Medial view

Lateral view

Tiktaalik
rosae

Sarcopterygian.
Shallow-water fish, but with:
Robust pectoral limbs, functional wrist
Well-supported ribs
Functional neck

Pentadactyl forelimb structure:

Hallux

Number of digits in
early
l amphibians
hibi
initially varied
from 4
4--13

Digits reduced in
living amphibians

Theropods

Convergent evolution
of external forms
using different
combinations of
skeletal elements

Historical constraints on adaptation

Ex:
E Giant
Gi
pandas
d thumb:
h b

Pandas related to bears and raccoons

(Carnivora).
Bears /raccoons are the most omnivorous
carnivores.
Use front p
paws to manipulate
p
food.

Giant panda are more dexterous than bears

and raccoons.
Diet restricted to bamboo.
Strip leaves between flexible thumb and
other digits.

Problems:

Primates have opposable thumbs, not

carnivores.
5 (not 4) remaining
g digits.
g

Constraints on adaptation
Pandas
P d thumb:
th b
Enlarged carpal bone:
Bears: slightly enlarged radial
sesamoid bone.
Carnivores: two 1st-digit abductor
muscles
muscles.
In bears, one muscle attaches
instead to radial sesamoid.
Abductor
Abd t and
d adductor
dd t muscles
l are
modified 1st-digit abductors.

Pandas thumb is modification of

carpal structures.

Limb
configurations
Amphibian and
reptilian
condition
(2 effective
ff ti
limb segments)
Mammalian and
archosaur
condition
diti
(3 effective
limb segments)
g

Muscular
Muscular sling
sling
of mammals

Elements of appendicular skeleton:

hi dli b
hindlimb
Pelvic girdle

Hindlimb

Mammals
Anterior

Posterior

Anterior

Posterior

 Acanthostega :

One of first amphibians with well-defined

well defined digits (8)
on forelimbs, hindlimbs.
Limbs extended laterally, not ventrally.
Joints (elbow, wrist, knee, ankle) present but with
little mobility.

Sues foot
(Tyrannosaurus rex)

Theropod
feet (4 toes)

Utahraptor

Vulture

Hallux

H
Human
pelvis
l i

Human pelvis

Knee

Elbow
Wrist

Ankle

Models of limb evolution

Account
A
tf
for ttwo major
j ttransitions
iti
in chordate limbs:
Aquatic fin folds to fins.
Fin-fold theory.

Fins to tetrapod limbs.

Axial and basal models.

Cephalochordate
condition

Fin-fold
Fintheory
h

Fish condition

Fin-fold model

Models of limb evolution

Vertebrata
Sarcopteerygii

Chondrich
hthyes

Gnathostomata
Crossopteerygii

Teleostei

Osteichthyes

Dipnoi

Selachii

Spiny-rayyed fishes

Silversidees, killifishes

Cods, angglerfishes

Lanternfiishes

Lizardfish
hes

Dragonfisshes

Salmons, trouts

Minnows,, characins, catffishes

Herrings,, anchovies

Tarpons, eeels

Bonytonggues, knifefishess

Bowfin

Gars

Sturgeonss, paddlefishes

Bichirs

Tetrapods

Coelacantths

Lungfishees

Chimaeraas

Sharks, raays

Lampreyss

Hagfishess

B
Basal
l model
d l
Axial model

Axial model
Lungfish condition is plesiomorphic.
g
with
Axial bones homologous
posterior long bones + carpals +
p
+ phalanges.
p
g
metacarpals
Evidence: embryonic and gene
expression patterns of vertebrate
limb development.

inductive sequence

Basal model
Coelacanth (crossopterygian)
condition is plesiomorphic.
Long
L
bones
b
are homologous
h
l
with
ith basal
b
l
fin elements (heterochronic change).
Radials are homologous with carpals +
metacarpals + phalanges.
Supported by phylogenetic evidence
for monophyly
f
p y y of
f the sarcopterygians.
p
yg

Context of evolution of tetrapod limb

Late Silurian Devonian (410-400 mybp):

Diversification of plants into semi-aquatic and terrestrial

habitats.
Formation of wetlands and rivers with abundant plant
life.
Appearance of terrestrial insects.

Transition from fin to limb preceded that from aquatic

to terrestrial habitats.
Early limbs sufficient for shallow, muddy waters.
Multiple, clearly defined digits.
Weak girdles and limbs,
limbs weak wrists
wrists.

Tetrapods: reduction of dermal bone, elaboration of

endochondral elements.
elements

Structure of the vertebrate skeleton

Two major sections:

Axial skeleton
Appendicular skeleton

Heterotopic elements

Articulations

Some heterotopic elements

P i bacula
Penis
b
l

Structure of the vertebrate skeleton

Two major sections:

Axial
A i l skeleton
k l t
Appendicular skeleton

Heterotopic elements
Articulations (arthroses):

Synarthroses

Amphiarthroses

Diarthroses

(forming
meniscus
layers)

Femoral ballballand--socket
and
joint

Trends in the skeletal system

y
Internalization: from external dermal skeleton to
internal endochondral skeleton.
Differentiation of regions of the skull and axial
skeleton.
Cephalization:
C h li
i
independence
i d
d
of
fh
head
df
from
postcranial skeleton.
Development and specialization of pentadactyl
limbs.
Relocation of limbs from lateral to ventral.
Interlocking of vertebrae via zygapophyses.
Reduction (loss and fusion) of skeletal elements.