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Comparative Anatomy of the Vertebrates:

[ CHAPTER 10:

Girdles, Fins Limbs and Locomotion]

OUTLINE I. Pectoral Girdle a. Ostracoderms b. Teleost c. Chondrichthyes d. Early Tetrapods - clavicles
OUTLINE
I. Pectoral Girdle
a. Ostracoderms
b. Teleost
c. Chondrichthyes
d. Early Tetrapods
- clavicles
- coracoid plate
- coracoid process
- scapula
e. Mammals
II. Pelvic Girdle
-
pelvic plate
-
sacroiliac joint
-
prepubic cartilage
-
epipubic and hypoischial bone
-
innominate bone
-
relaxin
III. Fins
A. Fin Rays
B. Skeletal Base of Fin Rays
C. Paired Fins
D. Medial Fins
E. Caudal Fins
F. Origin of Paired Fins
IV. Tetrapod Limbs
A. Manus
B. Pes
C. Origin of Limbs
D. Locomotion on Land without Limbs

APPENDICULAR SKELETON girdles, fins and limbs

Limb

buds

where tetrapod limbs arise during

embryogenesis

functionless vestiges which bespeaks an ancestor with functional tetrapod limbs

or

may

be

transitory,

incomplete

Fin folds where fins arise during embryogenesis

Fin folds – where fins arise during embryogenesis I. PECTORAL GIRDLES – dermal + replacement bones

I. PECTORAL GIRDLES dermal + replacement bones

a.)

OSTRACODERMS

girdles remain unclear

with

pectoral

appendages;

b.) TELEOSTS cleithrum as the major bone of the girdle

no clavicle

scapulocoracoid (coracoid+scapula)

c.)

CHONDRICHTHYES

components

calcified

endoskeletal

d.) EARLY TETRAPODS – interclavicle (addn’l membrane bone)

no posttemporal (braced the girdle against the skull in fishes)

supracleithrum is missing

*Please refer to The Skeletal System 4 Handout for the tabulated components of pectoral girdle in representative vertebrates.

CLAVICLES fate is correlated with that of the coracoid

clavicle and/or coracoid and procoracoid brace the scapula against the sternum

long bones of furculum (wishbone) in birds

CORACOID PLATE 2 ossification centers:

a. Anterior gives rise to procoracoids

b. Posterior gives rise to coracoids

CORACOID PROCESS OF THE SCAPULA vestige of the procoracoids and coracoids in Eutheria

SCAPULA present in all tetrapods that retain any vestiges of anterior limbs

the glenoid fossa for

articulation of the girdle with the head of the humerus

bears part

or all

of

e.) MAMMALIAN SCAPULA (Recall scapula of cat!):

1. Scapular spine

2. Supraspinous fossa

3. Infraspinous fossa

4. Acromion process

Mammalian clavicle large in monotremes, insectivores and primates

in bats, brace the scapula against the sternum

no clavicle in cats (to withstand the shock of landing upright on their forelimbs), ungulates (facilitates grazing) and ungulates

Observation: Dermal bones predominate in the pectoral girdle of bony fishes, whereas replacement bones predominate in tetrapods.

Dermal bones predominate in the pectoral girdle of bony fishes, whereas replacement bones predominate in tetrapods.

Comparative Anatomy of the Vertebrates:

[ CHAPTER 10:

Girdles, Fins Limbs and Locomotion]

II. PELVIC GIRDLES no dermal bones

1. PELVIC (ISCHIOPUBIC) PLATES paired (brace for the pelvic fins) that meet in the midventral pelvic symphysis in fishes

fused as one structure in chondrichthyes and lungfishes

with 2 ossification centers to form a pubis and an ischium in tetrapods acetabulum for articulation of head of femur ilium from blastema dorsal to the pelvic plate pubic, ischial and ishiopubic symphyses absent in birds to provide a wider outlet from the pelvic cavity for laying eggs pelvis sacrum + girdle (amniotes); encircles caudal end of the coelom pelvic cavity contains the urogenital organs; terminal portion of large intestine

2. SACROILIAC JOINT junction b/n sacrum and ilium; shock absorber; immobile

3. PREPUBIC (YPSILOID) CARTILAGE urodeles; for attachment of respiratory muscles

4. EPIPUBIC AND HYPOISCHIAL BONE reptiles, monotremes and marsupials; epipubic bone is marsupial bone (supports marsupium) in marsupials

Archaeopteryx no synsacrum

5. INNOMINATE (COXAL) BONE ankylosed ilium, ischium and pubis in mammals

6. RELAXIN ovarian hormone which softens the fibrocartilage of pelvic symphyses during parturition

2. LEPIDOTRICHIA bony; teleosts

Actinotrichia distal to the fin rays; stiffen the fin rays

B. SKELETAL BASE OF FIN RAYS

1. BASALIA absent in specialized teleosts

2. RADIALIA vestiges in specialized teleosts

Archipterygium ancestral fin type

C. PAIRED FINS

1. LOBED FINS sarcopterygians (actinistians and rhipidistians); fleshy proximal lobe and membranous distal portion

2. FIN FOLD FINS chondrichthyans; with broad base; basalia: propterygia, mesopterygia, metapterygia

3. RAY FINS actinopterygians (most like tuna have no pelvic fins for streamlinings); very flexible (teleosts)

4. SPINY FINS acanthodians; supported by hollow spines (not fin rays)

Biserial fin with two series of radials; e.g. dipnoan Neoceratodus

Claspers modified basalia in male chondrichthyans

D. MEDIAN FINS

1. DORSAL FINS act as keels, keeping motionless fishes from rolling to the left and right; may be used for locomotion in rare cases; impt in Rajiformes (inc. number), lampreys and bony eels (elongated)

2. ANAL FINS modified as gonopodia

(analogous to claspers) in some viviparous teleosts

E. CAUDAL FINS

1. HETEROCERCAL notochord turns upward; placoderms, Paleozoic and modern sharks, some acanthodians, and chondrosteans (sturgeons and spoonbills)

 

2. HYPOCERCAL vertebral column turns downward; ichthyosaurs (result of evolutionary convergence)

 

III. FINS

Functions:

3. DIPHYCERCAL externally symmetrical, vertebral column ends with a little upbending; dipnoans and Latimeria

1. steering devices for changing direction

2. stabilizers to prevent the body from rolling

3. control of body inclination when swimming away

from the horizontal

4. serve as brakes to decrease forward motion

A. FIN RAYS

4. HOMOCERCAL notochord turns far dorsad; teleosts

*Heterocercal condition most primitive; the rest are modifications of it

Comparative Anatomy of the Vertebrates:

[ CHAPTER 10:

Girdles, Fins Limbs and Locomotion]

*Similaities in morphologic features of caudal fins may be the result of convergent evolution

F. ORIGIN OF PAIRED FINS

formula);

therapsids and mammals with pentadactyl limbs

late

2-3-3-3-3

in

1.

FIN FOLD HYPOTHESIS paired fins are

derived from a pair of continuous fleshy folds of lateral body wall analogous to the

PREPOLLEX elongated carpal; can be seen in a panda

metapleural folds of an amphioxus; no evidence; of historical interest only

POLLEX thumb

2.

GILL ARCH HYPOTHESIS pectoral and pelvic

girdles are modified gill arches, and the

MODIFICATIONS OF MANUS:

skeleton within the fin is an expansion of the gill rays

1. reduction in the number of bones by evolutionary loss or fusion

3.

FIN SPINE HYPOTHESIS membranes in 2 pairs develop in rays supported by radial

2. disproportionate lengthening or shortening of some of the bones

elements

3. increase in the number of phalanges

*Pelvic fins appear to be a gnathostome feature.

*Spines in acanthodians derived feature

feature. * Spines in acanthodians – derived feature IV. TETRAPOD LIMBS * Early tetrapods – first

IV. TETRAPOD LIMBS

*Early tetrapods first segment is horizontal from the trunk; second segment perpendicular to the first, directed downward; among amphibians and basal lizards

THREE SEGMENTS:

1. PROPODIUM upper arm, thigh

2. EPIPODIUM forearm, shank

3. AUTOPODIUM manus, pes; most modifications

PATELLA (KNEECAP) sesamoid bone; in birds and mammals; protects the joint from the abrasive action of the tendon

PENTADACTYL LIMB five-digit limb

A. MANUS

1. CARPUS wrist with 3 rows of bones:

a. Proximal Carpals

radiale

intermediale

ulnare

pisiform sesamoid bone; in most reptiles and mammals

b. centralia 3 or more bones

c. distal carpals 5

hamate fused 4 th and 5 th distal carpals

2. METACARPUS skeleton of the palm

3. PHALANGES bones of the digits; 2-3-4-5-3 (generalized phalangeal

ADAPTATIONS OF MANUS:

1.

Flight

CARPOMETACARPUS fused three distal carpals and three metacarpals; in birds

ALULA first finger in brids; for braking

PATAGIUM wing membrane; pterosaurs, bats, gliding lemurs; a result of convergent evolution if present in unrelated mammals

2.

For life in the ocean became flippers

3.

For swift-footedness

PLANTIGRADE common in mammals with pentadactyl limbs (monotremes, marsupials, insectivores, bears, arboreal raccoons and primates); primitive tetrapod stance

DIGITIGRADE rabbits, rodents and most carnivores; increased speed and agility and more silent than plantigrade

UNGULIGRADE stance that is well- suited for running

a. PARAXONIC FEET body weight borne on two parallel axes; artiodactyls

b. MESAXONIC FEET body

weight borne on the middle digit; perissodactyls

4.

For grasping accomplished by flexing the fingers at each interphalangeal joint; primates, rodents; opposable thumb

B. PES

1. TARSALS wrist with 3 rows of bones:

a. proximal tarsals

fibulare

intermediale

tibiale

b. centralia 3 or more bones

Comparative Anatomy of the Vertebrates:

[ CHAPTER 10:

Girdles, Fins Limbs and Locomotion]

c. distal tarsals 5

cuboid fused 4 th and 5 th tarsals

2. METATARSALS skeleton of the palm

3. PHALANGES bones of the digits 2-3-4-5-4 (Sphenodon); 2-3-4-4-0 (alligator); 2-3-4-4-0 (turtles); 2-3-4-5-4 (generalized for reptiles); and 2-3-3-3-3 in early therapsids and mammals

PREHALLUX vestiges of a tarsal or metatarsal

HALLUX great toe

ASTRAGALOCALCANEUS fused proximal tarsals and a centrale; in lizards

TIBIOFIBULA fused of tibia and fibula (frogs); splinter in birds; lost in deer and other ungulates

TIBIOTARSUS fused of tibia and proximal tarsals (birds)

TARSOMETATARSUS

metatarsals

fused

distal

tarsals

and

*There is an intratarsal joint between the tibiotarsus and tarsometatarsus and a joint between the tarsometatarsus and toes.

ZYGODACTYLY X; woodpeckers and parrots

*Mammals have a hinge joint instead of an intratarsal joint. The tibiale is the principal weight-bearing bone of the ankle.

METATARSAL arch in hominoids; distributes the body weight over four solid bases; absorbs some of the shock generated by bipedal locomotion; provides “spring” for walking and running

*In wriggling seals, walruses, cetaceans and sirenians, the anterior flippers are for maneuvering. Wirggling seals move using their posterior flippers and with lateral undulations of the trunk. Walruses, cetaceans and sirenians move by dorsal and ventral undulations.

C. HYPOTHESES (ORIGIN OF THE LIMBS):

1. MODIFICATION OF EXISTING STRUCTURES (supported by fossil record)

Rhipidistian pectoral fin with basal bone articulates proximally with scapula and distally with a pair of radials

Loss of fin rays and modifications of distal radials could have produced the skeleton of the tetrapod limb

Preaxial and postaxial radials could have formed the digits

2. FORMATION OF NEW FEATURES (based on developmental studies)

Initial cell proliferations develop the limb buds at the sides of the trunk of the body

Second period of cell proliferation occurs at right angles to the limb axis

giving rise to the

(distal to the wrist) digits

D. LOCOMOTION ON LAND WITHOUT LIMBS

1. SERPENTINE or LATERAL UNDULATION by forming loops; limbless lizards and snakes

2. RECTILINEAR LOCOMOTION gliding forward on the substrate while keeping the entire body in a straight line; depends on generating friction between sections of the ventral skin and the substrate

3. SIDEWINDING rattlesnakes and snakes (desert)

4. CONCERTINA MOVEMENT modified serpentine movements; bracing S-shaped loops against the burrow wall and exerting horizontal force while thrusting the head and forebody forward