Examine the vertebrae of a shark and a tuna (derived bony fish), as well as a mounted bony fish skeleton.

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1 EEB 3273/Schwenk LAB 2-3: AXIAL AND APPENDICULAR SKELETON Preparation H & W, Chap. 5 (small print) H & W, Chap. 6 (small print; ignore bones of the hands & feet) Background Together, the axial (body axis or midline) and appendicular (bilaterally symmetrical girdles and limbs) skeletons compose the body, or postcranial skeleton (as opposed to the head skeleton). The postcranial skeleton comprises the vertebral column, ribs, the pectoral and pelvic girdles, and their associated fins and limbs. It serves as a storehouse for various minerals necessary for metabolism, protects internal organs, provides rigid structure for the body, and due to its segmented, hinged nature, combines with associated muscles for movement. In contrast to the rigid exoskeleton of arthropods, the internal skeleton is stronger and has allowed vertebrates to reach much larger body sizes while remaining relatively agile. The axial and appendicular skeletons are composed mostly of endochondral bone and cartilage. A few dermal bones are, however, associated with the pectoral girdle in some vertebrates. Today's Lab This lab is composed of three major parts. First, we examine the modifications seen in the axial skeleton relating to life in water (fishes) vs. life on land (tetrapods), comparing the latter to secondarily aquatic tetrapods (e.g., cetaceans). Second, we explore the morphology of the pectoral and pelvic girdles and appendages in fishes as compared to tetrapods. Third, we consider some of the modifications of the tetrapod body skeleton associated with four ecomorphological types: cursorial (running), fossorial (digging), scansorial (climbing), and volant (flying). The bone names for which you are responsible are listed in this handout. I. AXIAL SKELETON A. RELATIONSHIPS OF THE VERTEBRAL COLUMN 1) Fishes (H & W, Fig. 5-2) The vertebrae of fishes aid in the production of lateral undulations for swimming. The entire vertebral column is subjected to similar stresses and, therefore, shows little regional differentiation. It primarily resists compression caused during contraction of the segmented body musculature for swimming. Since fishes are buoyant in water, their axial skeleton does not need to resist the shear forces created by gravity. Two basic types of vertebrae can be identified: trunk (body) and caudal (tail). Material: Examine the vertebrae of a shark and a tuna (derived bony fish), as well as a mounted bony fish skeleton. Identify: trunk vertebrae caudal vertebrae centrum neural arch neural canal hemal arch hemal canal notochord (where present) ribs median fin tail fin EEB 3273, Lab 2, Page 1

2 2) Non-Mammalian Tetrapods (H & W, Fig. 5-4) As tetrapods moved onto land, the body skeleton became increasingly important for support and limbed locomotion. In response to gravitational stresses, tetrapods show more ossification of the body skeleton than most fishes and the development of intervertebral articulations (pzygapophyses + others in some groups) that help to interlock to stiffen the column and prevent shear. The vertebral column also shows further regional differentiation/specialization into cervical (neck), trunk, sacral (fused vertebrae where the pelvic girdle attaches) and caudal vertebrae. Material: Consider how the transition to terrestrial life has reshaped the axial skeleton in amphibians (a frog, Rana, and a salamander, Necturus) and reptiles (lizard, alligator and bird skeletons). Identify: cervical vertebrae trunk vertebrae sacral vertebrae/sacrum caudal vertebrae centrum neural arch neural canal neural spine/process atlas c.1 axis c.2 (not specialized) prezygapophyses postzygapophyses transverse processes hemal arch (only in caudal vert. of crocs) ribs uncinate process (birds) sternum 3) Mammals (H & W, Fig. 5-5, 5-6, 5-7) Mammals show the greatest degree of regionalization in the vertebral column. The vertebral column is vital not only for locomotion and protection of the spinal cord, but it also serves as an integral part of diaphragmatic respiration. The vertebral column is differentiated into cervical, thoracic, lumbar, sacral, and caudal regions (thoracic + lumbar = trunk). Be sure to be able to identify vertebrae from the five different spinal regions in mammals, as well as the terms listed below. Material: Examine an articulated cat skeleton, the isolated vertebral column of a horse (be sure to orient this correctly), whale vertebrae, and the cervical vertebrae of another small cetacean (dolphin), plus any other material present. Identify: cervical vertebrae thoracic vertebrae lumbar vertebrae sacral vertebrae/sacrum caudal vertebrae centrum neural arch neural canal neural spine/process atlas c.1 axis c.2 dens/odontoid process prezygapophyses postzygapophyses pleurapophyses transverse processes intervertebral disc (may not be preserved) ribs sternum Zygopophysis Hint: postzygapophyses feel bad about being stuck in back so they are down-and-out; prezygapophyses are happy about being in front and are therefore up-and-in (in reference to the direction that the articular facets the smooth surfaces where they make contact face) EEB 3273, Lab 2, Page 2

3 B. RIBS AND STERNUM (H & W, Fig. 5-5, 5-6, 5-7, 5-8) In terrestrial (tetrapod) vertebrates, ribs serve a much more important role in support and locomotion than in fishes. They protect the viscera and prevent the trunk from compressing when the body is lying on the ground. Furthermore, in amniotes (reptiles and mammals) the ribs have become vital to the expansion of the thoracic cavity during breathing. As a result, the ribs of birds and mammals show various modifications that allow for the insertion of intercostal (between rib) muscles. Some ribs have costal cartilages that connect them ventrally to the sternum (part of the axial skeleton in the ventral midline). This connection forms a strong but flexible box which is helpful in absorbing the intense shock produced in the pectoral girdle by running quadrupeds as the forelimbs strike the ground. In some climbing and digging mammals, some ribs become flattened blades that nearly overlap, or even fuse together, creating a more rigid thorax that resists dorso-ventral bending (e.g., anteaters). Birds have uncinate processes on their ribs that extend posteriorly, overlapping rib behind. This also helps to stiffen the rib cage to resist the stresses of flight. In most vertebrates, ribs are associated with nearly all vertebrae, including the cervicals (with the exception of c.1 and c.2, the atlas and axis used for head movement). However, a diagnostic feature of mammals is that there are no ribs on all 7 cervical vertebrae (monotremes have fused ribs on c. 3-7). This is the principal way to distinguish a cervical from a thoracic vertebra in a mammal there are rib facets on thoracic vertebrae and no rib facets on cervical vertebrae. Nearly all mammals have the same number of cervical vertebrae (7). This includes giraffes with very long necks and cetaceans with virtually no neck (see cetacean cervical vertebrae specimen). The constancy and apparent evolutionary stability of this number is a well-known example of an evolutionary constraint in other words, there is something that seems to prevent an evolutionary change (+/-) in this number. Recent work suggests that the constraint arises from high cancer rates associated with mutations that create more or fewer cervical vertebrae. Nevertheless, there are a few species that have either 6, 8 or 9. Can you imagine which mammals these might be and why/how they seem to have circumvented the constraint? II. PECTORAL GIRDLE, PELVIC GIRDLE, AND APPENDAGES 1) Fishes (H & W, pp , Figs. 6-1, 6-2) Material: (preserved shark skeletons, bony fish skeletons) Compare the pelvic and pectoral girdles of the shark and the actinopterygian fishes, Amia and Perca. Note especially that the girdles of the shark are composed of a single cartilage while those of the bony fishes are paired, with several bones composing each half. Compare also the breadth of connection between the fin radials and the girdles. In what ways would these differences affect the mobility of the fins? Identify: PECTORAL GIRDLE PELVIC GIRDLE PECT. AND PELVIC FINS Dermal Bones cleithrum Endochondral Bones coracoid scapula/scapular process puboischiadic bar iliac process EEB 3273, Lab 2, Page 3 basal pterygiophores radial pterygiophores fin rays: lepidotrichia in bony fishes (dermal bone) ceratotrichia in sharks (keratinous)

4 2) Non-Mammalian Tetrapod Condition (H & W, pp , Figs. 6-5, 6-6, 6-7) Material: salamander (Necturus), lizard, alligator and bird skeletons Amphibians and many living reptiles retain ancestral features of both the pectoral and pelvic girdles. Note the partial ossification of the girdles in Necturus versus the heavy ossification seen in reptiles. Why might this be? In birds, both girdles are highly modified for flight. The furcula ( wishbone ) is a new element in the pectoral girdle representing the fusion of the dermal clavicles and probably the interclavicle, as well. What is the function of the furcula? Identify: PECTORAL GIRDLE scapula (endochondral) coracoid (endochondral) clavicle (dermal) interclavicle (dermal) FORELIMB humerus ulna radius carpals (wrist) metacarpals (palm) phalanges (digits) PELVIC GIRDLE ilium ischium pubis synsacrum (birds) 1 HIND LIMB femur tibia fibula tarsals 2 (ankle) metatarsals (instep/foot) phalanges (digits) HIND LIMB OF BIRDS femur tibiotarsus 3 tarsometatarsus 4 1 The synsacrum of birds consists of the sacrum + additional fused vertebrae from the trunk and tail. In addition, some or most of the pelvic girdle bones are fused to the synsacrum, creating a very rigid structure to resist the stresses of flight, landing and take-off 2 The calcaneus or heel bone is a modified tarsal) 3 The tibiotarsus is a fusion of the tibia and some of the proximal tarsal bones 4 The tarsometatarsus is a fusion of the distal tarsal bones and the metatarsals. Like the cannon bone of some hoofed mammals, it adds an additional segment to the limb 3) Mammals Monotremes (Handout; Figs. 6-5, 6-6) Material: monotreme skeleton (echidna/spiny anteater, Tachyglossus sp.) In the synapsid stem lineage leading to mammals, a new endochondral bone developed in the pectoral girdle just posterior to the existing coracoid. Unfortunately, this bone is also called a coracoid, specifically, the posterior coracoid. Among living mammals, this condition is seen only in the monotremes, which retain both coracoids, the "new" posterior coracoid and the "old" anterior coracoid. In addition, late synapsids evolved new bones projecting anteriorly from the pelvic girdle called the epipubic bones. These are retained in monotremes and marsupials and are even found in extinct, fossil members of the eutherian mammals. These rod-like bones serve as attachment sites for muscles and tendons that help to stiffen the abdomen during locomotion (you might read that these bones serve to support the marsupium, or pouch, but this is no longer thought to be the case). In non-mammalian bony vertebrates, the pelvic girdle on each side consists of three, separate bones: a dorsal ilium, an anterior pubis and a posterior ischium. In mammals, however, these three ossifications EEB 3273, Lab 2, Page 4

5 fuse into a single large bone called the innominate bone. Although a single bone in adults, in embryos and juveniles one can observe the original, three separate bones. The three bones meet in the acetabulum the socket for the femur. The two pubic bones meet anteriorly in the midline where they are tightly fused with a fibrous cartilage joint called the pubic symphysis. The iliac crest (the ilia bones) on each side attaches to the sacrum forming the sacro-iliac joint also a tight, fibrocartilage joint. Identify: PECTORAL GIRDLE/LIMB scapula anterior coracoid posterior coracoid clavicle interclavicle humerus ulna radius carpals metacarpals phalanges PELVIC GIRDLE/LIMB ilium ischium pubis innominate bone epipubic bone acetabulum femur tibia fibula tarsals metatarsals phalanges 4) Mammals Therians (Marsupials and Eutherians) (Figs. 6-11, 6-12, 6-13, 6-16, 6-17, 6-18) Material: marsupial and eutherian mammal skeletons/limbs and girdls In marsupial and eutherian mammals, the pectoral girdle is highly simplified: only a single element remains, the large endochondral scapula. The ancestral anterior coracoid is lost and the "new" posterior coracoid remains only as the coracoid process, now fused to the scapula. The dermal bones are lost except for the clavicle, and even this is reduced or lost in many cursorial (running) mammals. The pelvic girdle of The epipubic bones of the monotreme/marsupial pelvic girdle are lost in eutherian mammals. Identify: PECTORAL GIRDLE/LIMB PELVIC GIRDLE/LIMB scapula coracoid process scapular spine clavicle humerus ulna radius carpals metacarpals/cannon bone* phalanges ilium ischium pubis acetabulum femur tibia fibula epipubic bone (marsupials only) tarsals metatarsals/cannon bone* phalanges *in some hoofed animals the metacarpals and metatarsals are reduced in number and fused into a cannon bone that forms an additional, vertical element of the leg, with the animal walking on the nails of its toe tips or hooves) EEB 3273, Lab 2, Page 5

6 DERIVED BONY FISH (teleost) SKELETON BONY FISH (basal/primitive) NOTE THE CONTINUOUS NOTOCHORD PRESENT EVEN IN THIS AND SOME OTHER BONY FISHES The vertebrae ossify around the notochord, but do not replace it during embryonic development. BONY FISH (derived) These are trunk vertebrae with attached ribs, therefore there are no hemal arches. Note that most derived bony fishes (teleosts) have two sets of ribs dorsal and ventral. In this species the dorsal ribs are tiny. Regardless, it is the dorsal, not the ventral ribs, that are homologous with tetrapod ribs.

7 1 = PUBOISCHIADIC BAR 2 = BASAL PTERYGIOPHORE 3. Claspers are found in males only; used as intromittent organ to introduce sperm into female cloaca 4 = RADIAL PTERYGIOPHORES; beyond these would be the CERATOTRICHIA that support the rest of the fin 1-2 = SCAPULOCORACOID BAR 3-7 = BASAL PTERYGIOPHORES 8 = RADIAL PTERYGIOPHORES (CERATOTRICHIA not shown) DERMAL BONES: pt = post-temporal bone of skull s.cl = supracleithrum cl = CLEITHRUM p.cl. = post-cleithrum ENDOCHONDRAL BONES: c = CORACOID sc. = SCAPULA br.c. = BASAL PTERYGIOPHORES (BASALS) BONY FISH (TELEOST) LEFT PECTORAL GIRDLE (lateral view)

8

9 BIRD PECTORAL GIRDLES (NOTE: fused clavicles + interclavicle = FURCULA or wishbone ) 1 = scapula 2 = coracoid 3 = interclavicle 4 = glenoid fossa (socket for humerus) BUDGIE (PARROT) note curvature of cervical vertebrae at rest ALLIGATOR PECTORAL GIRDLE

10 LIZARD PECTORAL GIRDLE (ventral view) DERMAL BONES 1 = INTERCLAVICLE 2 = CLAVICLE ENDOCHONDRAL BONES 3 = SCAPULA 4 = CORACOID OTHER 6 = GLENOID FOSSA 7 = STERNUM 8-9 = RIBS BASIC (ANCESTRAL) TETRAPOD PECTORAL GIRDLE (front view) 1 = suprascapula 2 = SCAPULA 3 = GLENOID FOSSA 4 = CORACOID 5 = CLAVICLE 6 = INTERCLAVICLE DIFFERENT LIZARD PECTORAL GIRDLS (lateral view, anterior to left)) PLESIOSAUR (extinct marine reptile) PELVIC GIRDLE SHOWING BASIC TETRAPOD CONDITION top = dorsal view bottom = lateral view p = PUBIS isc = ISCHIUM il = ILIUM (these attach to sacral vertebrae dorsally) acet = ACETABULUM (= socket for femur; always found at junction of all 3 bones)

11 BIRD SYNSACRUM: dorsal (left) and ventral (right) views HORSE FORELIMB PASTERNS = DIGIT BONES (PHALANGES) CANNON BONE = FUSED METATARSALS FIBULA IS LOST OR FUSED WITH TIBIA BIRD HIND LIMB

12 LIZARD LIMBS TETRAPOD FORELIMB MODIFICATIONS NOTE THAT WING BONES ARE HOMOLOGOUS, BUT NOT ALL OTHER PARTS OF THE WINGS ARE (e.g., bat and pterosaur wing surfaces are stretched skin, whereas in birds they are made of feathers)

13 Transverse process Transverse process Tibia Fibula NUCHAL LIGAMENT IN UNTULATES

14 MONTOTREME PECTORAL GIRDLE (platypus) HUMAN (EUTHERIAN) PECTORAL GIRDLE (scapula with coracoid process) + clavicle (not shown) INNOMINATE BONE (half pelvis) OF DOG (Eutherian) ADULT (bones fused) AND JUVENINLE (bones unfused)

15 A and B EXTINCT SYNAPSIDS; C = MARSUPIAL; D = CAT/EUTHERIAN (left lateral view; anterior to left) HUMAN EMBRYO UNFUSED BONES OF PELVIS FEMALE vs. MALE HUMAN PELVIS (not all pelves fit into this neat dichotomy)

16 A and B EXTINCT SYNAPSIDS; C = MARSUPIAL; D = CAT/EUTHERIAN (left lateral view; anterior to left) HUMAN EMBRYO UNFUSED BONES OF PELVIS FEMALE vs. MALE HUMAN PELVIS (not all pelves fit into this neat dichotomy)

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