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Chapter 10: Girdles, Fins, Limbs, Locomotion

Appendicular Skeleton
pectoral girdle
pelvic girdle
skeleton of fins and limbs

Girdles- brace fins and limbs against the counterforces that appendages transmit from the water or
from a substrate
- braced against one or more components of the axial skeleton

Agnathans, moray eels, caecilians, snakes, some lizards, amphisbaenians (except one genus): no paired appendages

During embryogenesis arise as

Tetrapod limbs limb buds
Fins fin folds

Pectoral Girdles
- skeletal complex in the body wall immediately behind the head that articulates with the anterior fins or limbs
Polypterus scapula and coracoid receive the forces transmitted to
the trunk from the fins, the posttemporal braces the
girdle against the caudal angles of the skull, and the
clavicle is braced against the opposite clavicle in the
midventral symphysis. (force of impact is diminished)
Teleost cleithrum has become the major bone of the pectoral
girdle; they have lost their clavicle; embryonic
endoskeletal coracoid unites with the scapula to form the
Cartilaginous fishes have only endoskeletal components and these do not
ossify; altho they become harden by calcification of the
cartilaginous matrix
Early tetrapods same with early bony fishes except addition of
interclavicle; and they have lost their posttemporal
Basal amphibian supracleithrum is missing; cleithrum tho present in early
amphibians, did not last long once tetrapods became
established on land

- Interclavicle of stem amphibians appears to have been persistent in lineages leading to amniotes
In alligators seen as an unpaired bone between the base of the two
In birds it is the unpaired cartilage or bone at the tip of the wishbone
In reptiles and monotremes some paired bones associated ventrally with the clavicles or

- Fate of clavicles: correlated with that of the coracoids

o Clavicles/ coracoids: likely to brace the scapula against the sternum and sometimes both do so
o Clavicles:
Urodeles and apodans missing

Nonavian reptiles other than lizards uncommon (altho vestiges develop transitorily in crocodilian
Birds long bones of the furculum
Mammals present

o Coracoids:
Tetrapods coracoids arise from an embryonic cartilaginous coracoid plate in
the lateral body wall extending ventrad from the glenoid region of
the scapula; anterior ossification centers in this plate give rise to
procoracoids; posterior centers give rise to coracoids
In eutheria neither procoracoid nor coracoid develop except for a vestige the
coracoid process of the scapula overhanging the glenoid fossa

o Coracoids and procoracoids: assist the clavicles or replace them functionally in bracing the scapula against the sternum
o Scapula: present in all tetrapods that retain any vestiges of anterior limbs because it bears part or all of the glenoid fossa for articulation of the girdle
with the head of the humerus
o in urodeles and anurans : Suprascapular ossification centers usually coalesce with scapular centers to become a single bone but suprascapula
remains independent in urodeles and anurans
- mammalian pectoral girdle: result of evolutionary changes that gave rise to early therapsids and their mammalian descendants
therapsid and monotremes Consist of interclavicle, clavicle, precoracoid, coracoid, and
eutherian mammals scapula and clavicle only (vestige of coracoid persists as a
coracoid process of the scapula)
humans coracoid process usually remains unattached to the scapula
until about 15yrs of age

o lateral aspect of the mammalian scapula

divided by a scapular spine into supraspinous and infraspinous fossae
these fossae are the anatomic origins of the strong muscles that insert on the humerus
scapular spine: insertion site for some of the appendicular muscles that arise on the vertebral column
acromion process: for muscular attachment; forms near the glenoid fossa

o mammalian clavicle
in monotremes, insectivores, primates large
in bats Strong and brace the scapula against the sternum
in felines greatly reduced into a mere splinter
enables cat to withstand the shock of anding upright on
their forelimbs after a leap because there are no rigid
connections between the scapulae, which receive the
initial impact, and any other part of the skeleton
in cetaceans and ungulates completely lost
absence in ungulates facilitates grazing

*dermal bones predominate in the pectoral girdle of bony fishes, whereas replacement bones predominate in tetrapods
- urodeles fail to develop any dermal bones in their girdles (or anywhere else in the body except skull)
- tetrapods almost never brace their pectoral girdles against the skull or vertebral column (said to have happened in a few pterosaurs)

Pelvic Girdles
- no dermal bones in the pelvic girdles of either fishes or tetrapods
Most fishes consist of a pair of simple cartilaginous or bony pelvic (ischiopubic)
plates that meet in a midventral pelvic symphysis and provide a
brace for the pelvic fins
Cartilaginous fishes and lungfishes two embryonic cartilages unite to form one adult plate
Teleost w/ short trunk pelvic plate lies immediately behind, or sometimes below, the
pectoral girdle and is often attached to it
Tetrapod embryos Develop cartilaginous pelvic plates
- each plate ossifies at two centers to form a pubis and a
more posterior ischium
Urodeles Pelvic plate may remain cartilaginous except for a small ischial
ossification center
- ilium: from an additional blastema dorsal to the pelvic plate
- acetabulum: accommodates the head of the femur at the junction of the pubis, ischium, and ilium
Tetrapod ilium braced against stout transverse processes of the sacral vertebrae (1 in amphibians, 2 in nonavian reptiles, more in birds and mammals)
Ribs often become ankylosed to the transverse processes, in which case the ribs become unrecoginizable except in embryos

(except in birds) there is a symphysis between either the two pubic bones( pubic symphyis), the two ischia (ischial symphysis), or both.
Symphysis- in the midventral body wall immediately anterior to the cloaca or its derivatives

Skeletal framework: forces transmitted to the two acetabula as a result of gravity or locomotion is distributed in two directions: to the vertebral column dorsally via ilia and to
the symphysis ventrally via ischial and pubic bones
- depends on the posture of the animal
Joint between head of femur and the girdle stabilized by the joint capsule and by muscles that approach the femur from opposing directions

Amniotes: sacrum and girdle form a bony enclosure, the pelvis, which encircles the caudal end of the coelom. Results in the formation of the pelvic cavity which contains the
urogenital organs and the terminal portion of the large intestine

Posture and mode of locomotion correlated with the shape, anatomic relationships, and proportional size of the ilium, ischium, and pubis

Frogs: ilia are slender and greatly elongated; extend from the sacral vertebra to the end of the urostyle, where they meet the ischia and pubic bones and where the
acetabulum is located
The joint between ilium and sacral vertebra is free to move and acts as a shock absorber
Tetrapods: sacroiliac joint is not freely movable

- weaker posterior limbs that can scarcely lift the sagging belly of a substrate w/o the buoying effect of water
- pelvic girdle is subjected to little stress
- pelvic plate differs little from that of fishes other than being braced against a single vertebra by a weak ilium
- usually lose their gills and develop lungs
- a slim median prepubic (ypsiloid) cartilage extends from the girdle forward in the linea alba for two or three somites: respiration
- structure of pelvic girdles is coreelated with their body structure and mode of locomotion
- ilium: braced against additional vertebra
- pubis: directed away from the ischium, resulting in a triradiate girdle
- ornithischian dinosaurs and birds: ischium and pubis are parallel and directed caudad
- a broad ischiopubic fenestra develops between ischium and the pubis on each side in some reptiles
- in alligators: opening in the wall of the acetabulum results from the incompletion of the wall of the socket

epipubic and hypoischial bone develop in association with the pelvic girdle of reptiles
- one or both are present also in monotremes and marsupials
- epipubic bone or marsupial bone in marsupials coz it supports the marsupial pouch

Modern birds:
- enormously expanded ilia and ischia and united with the synsacrum
- such girdles provide broad surfaces for attachment of hind limb muscles used in bipedal stance and for the thrust required by some species for flight takeoff
- no ischial or pubic symphysis for laying eggs; pubic bones reduced to long splinters
Archeopteryx pelvic girdle: braced against 4 sacral vertebrae; no synsacrum

- ilium, ischium, and pubis ankylose early in postanal life to form a left and right innominate (coxal) bone
- sacroiliac joint: where ilium on each side ankylose with the sacrum (dorsally)
- pubic or ischiopubic symphysis: where pubis and ischia meet to complete the walls of the pelvic cavity (verntroposteriorly)
- pelvic outlet: provides access to the exterior; where mammalian youngs are delivered
- relaxin: softens the fibrocartilage seprarating the bones at the symphysis during late pregnancy

- serve as steering devices for changing directions
- as stabilizers that prevent the body from rolling or wobbling
- as devices that control inclination (slope) of the body in swimming

- paired fins: serve as brakes that slow or halt forward motion

- pectoral fins: for forward locomotion
- posterior part of trunk, tail, caudal fin: forward thrust

- consist mostly of two surfaces of skin back to back

- stiffened by flexible rays (in the dermis) that radiate from a skeletal base
o lepidotrichia: in bony fishes
consist of jointed bony dermal scales aligned end to end
o ceratotrichia: in cartilaginous fishes
long horny rays composed of a material similar to that comprising the dorsal spines of sharks
o actinotrichia short and delicate; develops distally in both lepido and ceratotrichia
- the skeletal base from which fin rays of generalized fishes radiate, whether paired fins or median fins, consists of a row of cartilaginous or bony basalia and one or
more rows of radialia
chondrostean Polypterus (generalized ray finned) pectoral fins have three large basals and two distal rows of radials
Most specialized teleosts Have lost all basalia and retain mere vestiges of distal radials
Dipnoans and other rhipidistians Exhibit a wide variety of bases in both paired and dorsal fins
Achipterygium ancestral fin type

Paired fins:
- pectoral fins are braced against the glenoid fossa or glenoid region of the pectoral girdle
- pelvic fins are braced against a prominence on the lateral or posterior aspect of the pelvic plate
- 3 categories of paired fins:
1. Lobed fins
- consist of a fleshy proximal lobe containing the fin skeleton and its attached muscles and a membranous distal portion that is stiffened by fin rays
- dipnoan Neoceratodus: consists of a jointed central axis, the bones of which function like basals, and a series of pre and postaxial radials
- such a fin is said to be biserial because of the two series of radials
2. Fin fold fins
- have a broad base (broad in modern sharks)
- pro, meso, metapteryigia
- male chondricthyes: claspers- an intromittent organ which is a result of the modification of the basalia of the pelvic fin
3. Ray fins
- teleost: have lost components of the basal skeleton in time, but the fins have become increasingly flexible
- most ray finned fishes (eg Tuna): no pelvic fins
- lost of pelvic fins may be related to streamlining
- Characins: voracious teleosts; get an initial thrust out of the water with the caudal fin and, beating winglike pectorals, fly several yards using
appendicular muscles that, for a fish, are exceptionally large
Median Fins:
- fishes have one, two, or a series of median dorsal fins, and many have a midventral anal fin just behind the anus or vent
- acts as keels, keeping motionless fishes from rolling to the left and right
- may be used for locomotion in rare cases
- minimize deviation from a straight line that inevitably results from side to side thrusts of the locomotor tail
- Rajiformes: no anal fin; two dorsal fins far back on the tail
- Lampreys and bony eels: long dorsal fins which compensates for the absence of paired fins

Caudal Fins:
- classified based on their shape and the direction taken by the terminal portion of the notochord and vertebral column
Heterocercal Tail that contains dorsal and ventral lobes Placoderms, Paleozoic sharks, some
and in which the notochord turns upward acanthodians, modern sharks, 2 relict
into a large dorsal lobe chondrosteans (sturgeons and spoonbills)
Hypocercal Rare condition in which vertebral column Ichthyosaurs
turns downward
Diphycercal Vertebral column ends with very little Dipnoans, Latimeria
Homocercal Encased within a bony sheath, or urostyle Teleosts
turns far dorsad - some specialized teleosts have only two
hypurals and no epurals
von Baers theorem: we conclude modern caudal fins are modifications of a heterocercal condition

Origin of Paired Fins:

1. fin fold hypothesis: paired fins are derived from a pair of continuous flashy folds of lateral body wall analogous to the metapleural folds of an amphioxus
2. gill arch hypothesis of C Gegenbaur: pectoral and pelvic girdles are modified gill arches, and the skeleton within the fin is an expansion of the gill rays
3. fin spine hypothesis: in time acanthodians tended to lose all sines except the two pairs containing the fin rays

Tetrapod Limbs
- typically four limbs but some have lost one or both pairs; others modified into wings or paddles
Early tetrapods Short; first segment extended nearly
horizontal from the trunk; second segment Functions:
(Persists in urodeles, tortoises, basal was perpendicular to the first, directed Excellent shock absorbers
lizards) downward Support for body suspension
Other reptiles and mammals A rotation of the entire appendage toward the Greater leverage for locomotion on
body occurred; long axes of humerus and land
femur nearly parallel to vertebral column;
elbow directed caudad; knee cephalad

3 segments of the tetrapod limb skeleton:

1. Propodium
2. Epipodium
3. Autopodium
Positionally equivalent components of the anterior and posterior limbs
Name of Segment Skeleton
1. upper arm (brachium) Humerus
2. forearm (antebrachium) Radius and ulna
Anterior Limb 3. wrist (carpus) Carpals
4. palm (metacarpus) Metacarpals
5. digits Phalanges
Posterior Limb 1. thigh (femur) Femur
2. shank (crus) Tibia and Fibula
3. ankle (tarsus) Tarpals
4. instep (metatarsus) Metatarpals
5. digits Phalanges

Propodium and Epipodium

Humerus: bone of upper arm
o Similarity in tetrapods is more strking than the differences
o Variations in length, shape, and diameter adaptive modifications
o Mole: has expansions for insertion of massive shoulder muscles for digging
o Carinate birds: have a slender central cavity containing diverticula from the lungs
Radius and Ulna: bones of the forearm
o Radius
Former preaxial bone that ahs shifted in orientation
Articulates proximally with the humerus and distally with wrist bones on the thumb side of the hand
o Ulna
Longer, formerly postaxial bone
Articulates proximally with the humerus and radius, distally with the wrist bones on the side opposite the thumb
Sometimes fuses with radius or may be vestigials (frogs and bats)
Femur: bone of the thigh
Tibia and fibula: bone of the lower leg or shank
o May unite to form a tibiofibula (frogs)
o May be reduced to a splinter (birds)
Tibiotarsus tibia fuses with proximal row of tarsals
o May be lost (deer and other ungulates)

Patella or kneecap:
- a sesamoid bone develops in birds and mammals
- protects joints against abrasive actions
- ossifies in the tendon of insertion of the powerful extensor muscle of the thigh where the tendon passes over the complicated knee joint to insert on the tibia

Manus: the hand

wrist (carpus)
o consist of 3 more or less regular rows of carpal bones
proximal row
radial carpal/ radiale at the distal end of the radius
ulnar carpal/ ulnare at the end of ulna
intermedium between the two
*pisiform: in mammals and reptiles sesamoid bone at the ulnar end of the proximal row
middle row
Centralia often 3 or 4 in early tetrapods (2 in reptiles)
Distal row
5 distal carpals
o metacarpals: skeleton of the palm
o pentadactyl limb may have had as many distal carpals and metacarpalsas there were digits
o each digit consists of phalanges
o 2-3-4-5-3: early phalangeal formula for a pentadactyl hand
o 2-3-3-3-3: late therapsids and modern mammals with 5 fingers
Oldest known tetrapods 6-8 digits
Ichthyostega (hindlimb) 7 digits
Acanthostega (forelimb) 8 digits
Tulerpeton 6 digits
3 digits in Ichtyostega that occupy the position of the big toe in later tetrapods might be fulfilling the same ocomotor role as the big toe of a pentadactyl (5-digit) limb

Modifications of the manus:

- reduction in the number of bones by evolutionary loss or fusion
- disproportionate lengthening or shortening of some of the bones
- increase in the number of phalanges
Centralia: frequently unite with one of the proximal carpals or disappear
Most reptiles and mammals: have a single centralia and sometimes found among the proximal row of carpals
Hammate bone: result of the fusion of distal carpals 4 and 5

Phalanges or digits may be lost; corresponding metacarpal becomes vestigial or lost

Most amphibians 5 digits in the hind limb; 4 digits in the forelimb
Amphiuma 1-3 digits
Necturus lost metacarpals that corresponds to any lost ancestral digits
Labyrinthodont amphibians more number of wrist bones than modern amphibians

Embryonic intermidium and ulnare often coalesce during ontogenesis, an embryonic proximal carpal often coalesces with an adjacent carpal, and fusion between embryonic
centralia and proximal or distal carpals is common.

Prepollex a bone of unknown homology that develops in some species near the thumb or pollex

Hypothesis of Alberch and Gale: rather than a truncation of development (a paedomorphic loss of the last digit to form), a reduction in the number of primordial cells in the
developing limb bud causes reorganization within the limb resulting in digit reduction
Urodeles and Anurans (but only hands in anurans) - hands and feet dont generate locomotor thrust
- chiefly platforms, or podia, which provide friction while muscles
higher the limbs extend the legs
living nonavian reptiles and mammals (insectivores and primates) - hands tend to remain pentadactyl and to have 5 metacarpals and
a nearly full complement of carpals except central carpals or
crocodilians - the wrist have been reduced to five adult bones
birds - entire manus has been reduced
mammals - when centrale is present, it may lie in the distal row of carpals
(rabbits) or unite with the radiale and intermedium to form a
scapholunar bone of triple origin (cats)
- human fetuses: central until 3rd fetal month the fuses with radiale

Major Modifications of the Hand:

1. Adaptations for flight
a. In many
Marine amniotes (sea lions, cetaceans, sirenians, hand have become paddlelike flippers birds: hand
ichthyosaurs, plesiosaurs, seals, penguins) has an
Penguins - obtain thrust for swimming solely from their flipperlike wings aerodynamic
- webbed feet serve as rudders effect; loss
Sea turtles, fur seals, sea lions swim by rowing action of their long front flippers and fusion of
Wriggling seals (true seals), walruses, cetaceans, sirenians dont use flippers to produce locomotor thrust when swimming bones have
reduced the
hand to a rigid, tapering structure
i. Carpometacarpus: formed by the union of 3 distal carpals and 3metacarpals
ii. 3 fingers usually present # of phalanges reduced; covered by feathers
first finger: alula - elongated, prominent, and independently movable; for maneuver, alight, and take off
juvenile Opisthocomus claws present and used in climbing; lost in adults
songbirds short, broad wings; alula serves as an accessory airfoil
carnivorous birds long, broad wings for slow speed flight and quick landings
pterosaurs and bats hand is main part of the wing
4th finger of pterosaurs embedded in patagium (wing
bats: 5 fingers; normal thumb and other 4 elongated

lemurs patagium present but less developed

2. Adaptations for life in the ocean

a. Flippers generally flattened and stout; # of phalanges greatly increased
i. Ichthyosaurs 26 phalanges per digit
b. Some aquatic mammals have lost all traces of hind limbs

3. Adaptations for swift-footedness

a. Plantigrade - mammals with pentadactyl hands and feet; wrists, ankles, digits all rest on the ground; primitive tetrapod stance not associated with
b. Digitigrade mammals in which only first digit is reduced; bear weight on their digital arches with wrist and ankle elevated; rabbits, rodents, most
c. Unguligrade super reduced # of digits; walking on the tips of the remaining digits
i. Eohippus ad 4 digits on the manus
ii. Equus has a single digit
iii. Horses proximal row of carpals intact; distal row lacks only the 1 st carpal
iv. Artiodactyls weight of the body tended to be distributed equally between digits III and IV; thus arose the cloven hoof said to be
paraxonic because body weight is borne on two parallel axes; even number of digits
v. Perissodactyls body weight is increasingly tended to be borne on digit III (middle digit); said to be mexasonic foot odd number of digits

4. Adaptations for grasping

Many mammals able to flex their hand at the joint between palm and fingers
Rodents nibble food held between two hands; flexed in a manner which
faced one another
Primates wrapping the fingers around an object to held it securely using one
hand; accomplished by flexing the fingers at each interphalangeal
Humans opposable thumb formation of a saddle joint at the base of the
thumb where it meets the palm; strong adductor pollicis (thumb)

Pes: The Hind Foot

- comparable bone for bone with the manus except that there is no consistent equivalent to the pisiform bone
- Early tetrapods: 4 centralia in the ankle
- # of centralia become smaller as modes of locomotion expanded
- pes of amphibian: prehallux may be vestiges of a tarsal or metatarsal that was associated with an ancestral digit that has been lost
- anurans: smaller # of ankle bones than urodeles; tibiale and fibulare have elongated and united firmly at each end to articulate distally with the remaining tarsals in
an intratarsal joint (such an ankle provided with long, webbed metatarsals and phalanges are well adapted for pushing off a leap on land or swimming in water)
- living reptiles: display considerable loss and fusion of ankle bones (turtles to a lesser degree than others)
o astragalocalcaneus - reduced proximal row of tarsals to a single bone
o sphenodon : 2-3-4-5-4 (general formula for reptiles)
o alligators: 2-3-4-4-0
o turtles: 2-3-3-3-2
o birds: highly modified foot;
tibiotarsus- union of proximal tarsals and the lower end of the tibia
tarsometatarsus union of distal tarsals with upper end of three fused metatarsals
there is an intratarsal joint between the tibiotarsus and tarsometatarsus and a joint between the tarsometatarsus and toes
digitigrade stance; 4 toes and a few has 3; ostriches alone has 2
usually 3 toes are directed radially forward and one comes off the back of the foot, but in woodpeckers and parrots, two toes at the back
and the four forming an X (zygodactyly)
- mammals: lack an intratarsal joint but have a large hinge joint where the tibia and fibula meet the ankle
o tibiale- principal weight-bearing bone of the ankle
o fibulare- elongated backward in platingrade mammals; upward in digitigrade and unguligrades; heel bone of plantigrades
o hominoids: metatarsal arch or instep distributes the body weight on all four solid bases (the heel and ball of each foot)
o phalangeal formula: 2-3-3-3-3 (early therapsid and modern pentadactyl mammals and humans
o hallux or great toe opposable in many primates but not in humans; when the condition is present, it is correlacted with branchiation

Sculling and Galloping in Marine Mammals

- wriggling seals, walruses, cetaceans, sirenians: modes of swimming dictated by the size and shape of the body and the morphology of the limbs they have
wriggling, or earless, seals swims with their foreflippers adducted against their sides, their neck
retracted, and with lateral undulation of the posterior portion of their
trunks; tail is short and of no use in locomotion; posterior
flippersswept alternately to the left and right with each lateral
undulation cycle of the trunk propels pinnipeds forward
walruses swim by sculling also
cetaceans unlike seals, they derive forward thrust from a stiff, horizontal
bifluked tail, which is operated by alternate dorsal and ventral
undulations of the caudal portion of the trunk (a sort of legless

How pinnipeds maneuver on land

- fur seals, sea lions, walruses: adaptation that enables rhe hind flippers to change readily from a caudally directed swimming position to the equivalent of a tetrapod
stance; hind flippers are simply rotated beneath the body from a swimming position to a tetrapod position or the reverse
- wriggling seals: hind limbs permanently bound to the tail

Origin of Limbs
- paired fins of acient fish must have been the precursor of tetrapod limbs
- 2 hypotheses on the origin of the tetrapod limb center on:
o a modification of preexisting structures
o the formation of new features
*common to both hypotheses is the origin of the proximal elements of the limb
1. the limb axis extends through the radials of the fin with pre- and postaxial radials forming the digits
2. the development of the tetrapod limbs differs from that in ray-finned fishes
- tetrapod limb has a second period of cell proliferation at right angles to the limb axis distal to the wrist
o gives rise to digits
o suggest that digits are not a restricting of preexisting features
1. elongation of the bones of the epipodium
2. formation of hinge joints between the propodia and epipodia (now elbow and knee joints); and between the epipodium and wrist or ankle
3. rotation of the long axes of the humerus and femur to parallel the vertebral column
4. emergence of definitive manus and pes

Locomotion on land without limbs

Snakes - Modified vertebral column (400 or more flexible intervertebral joints), ribs, body wall musculature, skin
- Serpentine mode of locomotion/ lateral undulation: body should be metameric with respect to the adult
skeletomuscular system
- Rectilinear locomotion: gliding/ flowing 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
o Two sets of striated muscles are responsible for rectilinear locomotion:
A pair of slender costocutaneous muscles extends downward and backwad from high on each
rib to the dermis at the edges of each scute
A more powerful pair extends less obliquely from a scute to a lower end of a more caudal rib
- propelling in their burrow
o concertina movement: modified serpentine movement; bracing S-shaped lops against the burrow wall
and exerting horizontal force while thrusting the head and forebody forward, more caudal sections then

Amphisbaenians - use rectilinear progression but the entire skin moves, not just the ventral skin
- use a modified serpentine movement in their burrows
Rattlesnakes - sidewinding enables them to inhabit sandy desserts