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CHAPTER 1: INTRODUCTION COMPARATIVE VERTEBRATE ANATOMY y Descriptive morphology y Functional morphology y Ontogenesis y Phylogenesis PROTOCHORDATES y No vertebral column but

share vertebrate embryonic features o Notochord o Dorsal hollow central nervous system o Pharynx with paired pouches and clefts in the embryo stage o Kingdom Animalia Phylum Chordata Subphylum Urochordata Subphlyum Cephalochordata Subphylum Vertebrata (Craniata) *Chordates are animals with notochord in the embryo stage. Vertebrates are Chordates with Vertebrae VERTEBRATE BODY: GENERAL PLAN y All vertebrates share o Similar patterns of embryonic development o Primitive structural and developmental features y Some features altered by genetic change overtime results to anatomic diversity y Regional Differentiation o Head  Special sense organs for monitoring external environment  Brain - large enough to receive and process info. ; provide stimuli to body musculature  Jaws acquiring, retaining and macerating food  Gills in fishes for respiration  CEPHALIZATION larger braincases become movable independent of the trunk o Trunk  Coelom cavity houses viscera  Body wall muscle, vertebral column and ribs  Neck narrow extension of trunk; lacks coelom ; consists of vertebrae, muscles, spinal cords, nerves and elongated tubes ; connects head to trunk DUH! o Post anal tail  Commences at anus or vent hence post anal  Caudal continuation of body wall muscles, axial skeleton, nerves and blood vessels  Always present during embryonic stage  Amphibians have tails; some Aves have reduced tails called NUBBIN ; human embryos have vestigial post anal tail Adult humans have the COCCYX o Paired pectoral and pelvic appendages  Supported by internal skeleton  Operated by muscles  Some are vestigial have been completely lost BILATERAL SYMMETRY AND ANATOMIC PLANES y 3 principal body axes o Longitudinal  Superior above  Inferior below o Dorsoventral  Dorsal back  Ventral where mouth is located o Left-right axis  Terminates in identical structures at each end

METAMERISM y Serial repetition of body structures in the longitudinal axis y Seen in vertebrate embryos and is retained in most adults y Muscle segments are reflections of embryonic metamerism y Serial arrangement of o Vertebrate o Ribs o Spinal nerves o Embryonic kidney tubules o Segmental arteries and veins VERTEBRATE CHARACHTERISTICS: THE BIG FOUR 1. & 2. Notochord & Vertebral Column y Notochord o 1st skeletal structure to appear in embryo o At peak; rod of living cells located ventral to CNS and dorsal to alimentary canal o Extends from midbrain to tip of tail o Part near head incorporated to floor of skull except agnathans o Part in the trunk and tail surrounded by vertebrae o FATE IN ADULT VERTEBRAES  Fish, Urodeles and Primitive Lizards exists at length of trunk and tail; constricted w/in each centrum  Modern Reptiles, Aves lacks vestige as it develops  Mammals reduced to vestige w/in invertebral disks that separate successive centra ; vestige consists of pulpy nucleus, soft spherical mass of connective tissue.  Agnathans grows along w/ animal and paired lateral neural cartilages become perched on notochord lateral to spinal shord; cartilages are like neural arches but unknown if it is primitive, vestigial or entirely different o Notochord Sheath  When notochord persists as an important part od adult axial skeleton it turns in to a strong outer elastic and inner fibrous sheath y Vertebral column o Vertebrae - Cartilaginous or bony rigid support  Centrum deposited around notochord  Neural Arch forms over spinal cord  Various processes  Hemal arch near tai; surrounds caudal artery and veins

3. Pharyngeal Slits y Pharynx o Region of the alimentary canal showing pharyngeal pouches in the embryo y Pharyngeal Slits o Pouches rupture to the exterior to form slits o May be permanent, adult pharynx part of alimentary canal with slits adults that live in water and breathe through gills o May be temporary, adult pharynx is part of the alimentary canal connecting oral cavity and esophagus if the animal is going to live on land y Pharyngeal Pouches o Arises as diverticula of the pharyngeal endoderm o Invade the pharyngeal wall & grow towards the surface y Ectodermal Groove o Simultaneously grows w/ pharyngeal pouches o Grows toward each pouch y Branchial Plate o Separates groove from o When ruptured a pharyngeal slit is formed as a passage way between the pharyngeal lumen and the exterior y Pharyngeal Arches o Column of tissue that separates pharyngeal pouch or slit o Located between 2 successive pharyngeal pouches or slits, in front of the 1st and immediately behind the last o Covered b the integument and lined internally by endoderm o Lost when slits close or grooves disappear st o Contains 4 basic components ,found in front of 1 pouch and directly behind last pouch, or BLASTEMAS from which these components develop 1. Supportive skeletal elements 2. Striated muscles operate the arch 3. Branches of 5th, 7th, 9th or 10th cranial nerves innervate muscles and provide sensory input to CNS 4. Aortic arch connects ventral and dorsal aortas o Visceral skeleton skeleton of Pharyngeal arches o Branchiomeric muscle primitive musculature of gills o 1st or Mandibular arch upper and lower jaws and associated muscle, nerves and vessels o 2nd or Hyoid arch behind mandibular arch o Branchial arches arches that support gills y FATE IN ADULT VERTERATES o Permanent marine  Jawed marine vertebrates, primitive sharks 8 slits and pouches  Agnathans 15 pouches and slits  Urodeles retain 1-3 slits o Temporary terrestrial, aerial  Frog y Embryo 6 pouches y Tadpoles 4 gives rise to slits  Reptiles, Aves, Mammals y No gills develop in pouches and slits are transitory st y Chicks 5 pouches develop, 1 3 rupture to the exterior then close again y Mammals only 1-2 of many anterior pouches may rupture th y Cervical Fistula failure of cervical sinus (3rs and 4 slits) to close  Tetrapods y Pouches rarely gives rise to permanent slits st y 1 pouch auditory tube and middle ear cavity nd y 2 pouch of the palatine tonsil of mammals

y Walls of several pouches gives rise to endocrine tissue in all vertebrates y Pharyngeal arches are anatomic entities y Primitive vertebrate pharynx was for filter feeding out of respiratory water stream shifted to branchial then pulmonary respiration

4. Dorsal Hollow Central Nervous System y Arises as longitudinal NEURAL GROOVE in what will become the dorsal surface of the embryo y Groove closes over and sinks beneath the surface to become a hollow NEURAL TUBE dorsal to the notochord. Tube is wider anteriorly and this becomes the BRAIN with its VENTRICLES y CRANIAL & SPINAL NERVES connect CNS w/ various organs of the body ; metameric spinal nerves y PERIPHERAL NERVOUS SYSTEM is made up of nerves along with associated ganglia and plexuses y FATE IN ADULT VERTEBRATES o Cyclostomes, teleosts, ganoid fishes  Basic pattern of NEUROCOEL, cavity, instead of forming a groove the surface ectoderm dorsal to the notochord proliferates a wedge-shape NEURAL KEEL. Eventually keel separates from the surface and a cavity forms w/in it by rearrangement of the cells in its interior o Fishes and amphibians 10 cranial nerves o Reptiles, birds and mammals 12 cranial nerves (extra 2 are spinal nerves trapped w/in skull) VERTEBRATE CHARACTERISTICS: SOME SATELITE FEATURES y Features not unique to vertebrates o Bilateral symmetry o Metamerism o Presence of coelom feature of all trploblastic animals o Deuterosmes embryonic blastopore, initial opening of embryonic gut or archenteron becomes anus y Integument o Epidermis  Glands defensive, lubricatory, nutritive, pheromonal and homeostasis-maintaining  Terrestrial vertebrates have spines,scales,feathers,hair,claws and hooves  Cornifiesd shells layer of dead cells for protection from dehydration from exposure to air o Dermis  Bone provides heavy coat of armor and contributes to dense, sharp, pointed scales of shark, thin flexible scales of teleost y Respiratory mechanism o External respiration exchange bet. Animal and environment through highly vascularised membranes (gills) located on pharyngeal arches or derived from pharyngeal floor (lungs) o Some breath through skin or BUCCOPHARYNGEAL LINING o Embryos through extraembryonic membranes that lie just inside the porous egg shell or through lining of mother s uterus y Coelom o tube within tube coelom between body wall and digestive tract o Partitioned coelom in fishes, amphibians and some reptiles o PERICARDIAL CAVITY - houses heart o PLEUROPERITONEAL CAVITY houses other viscera, lungs o TRANSVERSE SEPTUM separates pericardial and pleuroperitoneal cavity o PLEURAL CAVITY houses lungs in reptiles, birds and mammals

TENDINOUS OBLIQUE SEPTUM partition in reptiles & aves TENDINOMUSCULAR DIAGPHRAGM - partition in mammals SCROTAL CAVITIES hoises testes PERITONEAL MEMBRANE encloses coelom PARIETAL PERITONEUM membrane lining the body wall VISCERAL PERITONEUM part lying on the viscera Parietal and visceral peritoneum both continuous via dorsal or ventral mesentries o Kidneys lie against dorsal body wall external to peritoneum RETROPERITONEAL y Digestive organs o Acquisition, temporary storage, processing and absorption of food and for elimination of unabsorbed residue o Oral cavity, pharynx, esophagus, stomach, intestine o Spiral valve intestine in sharks increases absorptive area w/out increasing body length o CECA or DIVERTICULA including LIVER, PANCREAS have special functions in digestive process o CLOACA chamber that receives urinary and reproductive ducts and opens to the exterior via VENT o ALL vertebrate embryos have cloaca but in teleost its so shallow ans mammals it is subdivided into 2 or 3 passageways y Urogenital organs o KIDNEYS & GONADS arise together in roof of coelom and share certain passageways o KIDNEYS / NEPHROI eliminates water; assist in maintaining electrolyte balance in blood  In primitive fishes fluid wastes accumulate in coelom then removed by kidney tubules  Man s complex kidney tubules collect fluids directly from blood capillaries  Tubules lead to pair of longitudinal ducts that open into cloaca or urinary bladder o GONADS ducts, accessory glands, storage chambers and copulatory mechanisms  All vertebrate embryos are bisexual w/ gonadal and duct primordial for both sexes  Female gonad primordial develops into ovaries  Male gonad primordial becomes testes o Cyclostomes lack reproductive ducts; sperm and eggs are shed into the coelom and exit via a urogenital papilla located immediately behind anus y Circulatory system o Blood consists of plasma, red and white blood cells and platelets confined in veins, arteries, capillaries and sinusoids o SINUSOIDS broad channels rather than tubes; prominent feature of invertebrate circulatory system more common in fishes than tetrapods o LYMPH VESSELS collect interstitial tissue fluids and conduct them to large veins o HEART forms ventral to embryonic pharynx; pumps blood forward to ventral aorta through aortic arches and into dorsal aorta  Fishes close to gills; single circuit heart; blood passes from heart to the gills wher o2 is acquired then to tissues  Tetrapods displaced caudad during development  Mammals loss of gills and reliance on lungs caused twocircuit hearts, where oxygenated blood in the heart is pumped to all parts of the body where O2 is released then blood low in O2 is pumped to the lungs then through the heart again o Branches from the anteriormost aortic arches carry blood to the head o o o o o o o

y Skeleton o Cartilage, bones and ligaments o Structural; for protection of visceral organs; site of attachment for muscles o Rigid because of mineralization of connective tissues o Consists of  Axial skeleton skull and vertebral column  Pharyngeal skeleton best developed in fishes; supports gills  Appendicular skeleton y Muscles o Trunk and tail muscles move axial skeleton o Budlike extensions from body wall muscles on fins or limbs move the appendicular skeleton o Muscles on pharyngeal arches move pharyngeal skeleton o STRIATED MUSCLES foregoing muscles VOLUNTARY o CARDIAC MUSCLES variety of striated tissue REFLEX o SMOOTH MUSCLES walls of hollow viscera REFLEX y Sense organs o EXTEROCEPTORS monitor the external environment  Mechanoreceptors  Chemoreceptors  Electroreceptors  Thermoreceptors  Receptors for radiation o PROPRIOCEPTORS - monitor activity of muscles, joints & tendons o VISCERAL RECEPTORS monitor the rest of internal environment

CHAPTER 2: CONCEPTS, PREMISES AND PIONEERS PHYLOGENY y Evolutionary history of a taxon y Relates taxon to an ancestral taxa in a continuous evolutionary line PHYLOGENESIS y Process that produces lineage; speciation formation of new species from existing ones y Requires more than a lifetime ONTOGENY y History of an individual ; from embryonic development to death including post-embryonic changes and aging ONTOGENESIS y Embryogenesis and subsequent changes ; genes y Requires a single lifetime VON BAERS LAW 1828 y Features common to all members of a major taxonomic group of animals develop earlier in ontogeny than do special features that distinguish subdivision of the group y Common embryonic features gradually modify accdg. To the adaptation of the species as it develops in its environment y BIOGENETIC LAW o Features that develop earliest in ontogeny are the oldest phylogenetically having been inherited from common ancestors and features that develop later in ontogeny are of more recent phylognetic origin TAXONOMY y Study of the principles and procedures employed in placing organisms in appropriate taxa in accordance to 1 or more predetermined criteria y Techniques of ordering taxa into hierarchies that reflect the criteria in a meaningful way y Criteria: o Phylogenetic genealogy of the group

o Morphological similarities and differences in phenotypes y Classification o Orderly arrangement of taxa that shows accurate interrelationships of simplest to most informative manner possible o Schemes based on similarity of structure, geologic record, rationality and intuition  Natural order or evolutionary taxonomists utilize criteria given, data gathered and relevant data from other disciplines compared  Cladistic classification for classifying fossils disregarding geneology; number of phenotypic similarities computed and plotted into a dichotomous branching diagram; considers phenotypic similarities insufficient because some species have similarities but have different ancestry  Genealogical or Family Trees  Visual representation of ancestry of a species  Do not show distribution in time, size of population or relaive number of species  Show lineages SYSTEMATICS y Arises from taxonomy and study of the evolutionary processes HOMOLOGY y HOMOLOGUE same organ in different animals under every variety of form and function y Structures in 2 diff. Species are homologous if the come from the same embryonic precursor o Middle ear of mammals and hyomandibular cartilage that suspends lower jaw from braincase of sharks  Same structure in different animals; dissimilar appearance and function o Precaval vein of cats and right common cardinal vein of lower vertebrates o Intermaxillary bone of human embryo and the premaxillary bone of adult apes o Humerus of crocodile, bird and mammal  Same appearance, location and function y SERIAL HOMOLOGY - serial structures like spinal nerves and vertebrae are homologous as a series wit hthe same series in another species ; individual components of the series are not homologues because they are not from the same embryonic tissue ANALOGY y Coincidental resemblances y ANALOGOUS - 2 structures have the same function but are not homologous o Jaw teeth of bony fishes and horny ectodermal teethe of jawless lampreys o Horns of cattle and rhinoceros o Thymus gland and bursa of Fabricius of birds ADAPTATION y Hereditary modification of a phenotype that increases probability of survival y Result of genetic changes visible in the phenotype NATURAL SELECTION y Charles Darwin ; elimination and perpetuation of weak and strong phenotypes y Genetic mutations and recombinations bring about varitations in phenotypes which renders a population of organisms more or less able to adapt to the environment y Leads to SPECIATION isolation of gene pool no longer needed y TELEOLOGICAL REASONING use of design, purpose or utility to explain a natural phenomenon CONVERGENT EVOLUTION

y Applied when 2 or more unrelated species occupying the same kind of environment concurrently or millions of years apart, develop a similar adaptive morphologic trait y Taxa are unrelated and approach one another in superficial resemblances as a result of adaptive mutations to similar environments o Streamlined body with dorsal fins, a bilobed tail and paddle like anterior limbs in ichthyosaurs, aquatic reptiles, and dolphins, aquatic mammals o Webbed feet in frogs, ducks and platypus o Wings in pterosaurs, birds and bats  Appendages are homologous, functions are analogous similar adaptations are a result of convergent evolution PARALLEL EVOLUTION y Development of similar structures in recently related but isolates taxa whose common ancestor lacked such traits y No opportunity for new mutations as they arose to be transmitted from one group to the other; isolated in a manner as to stop interbreeding y Taxa diverged from recent common ancestor then independently evolved certain similar traits that were no present in the ancestor o Australian marsupials resemble placental mammals like wolves, foxes, bears and cats to the extents of being indistinguishable; similar phenotypes in the 2 groups were not present in the postulated common therian ancestor; cause is unknown HOMOPLASY y Similar structural adaptations in different organism that are not attributed to homology or to common ancestry y Those cited in convergent and parallel evolution SPECIATION y Formation of new species preceded by geographical isolation of a population from other population of same species y Geographical isolation with genetic change is the matrix out of which new species evolve y Natural selection channelizes the evolving product in the direction of increasing adaptivenes y PHYLOGENESIS result of continuous speciation; manifests effect of evolution ORGANIC EVOLUTION y Plants and animals on earth have been changing aand the ones around us today are descendants of those which were here earlier y Life comes from pre-existing life y Natural explanation for the variation of species y theory of the mutability of species o species appeared coming from no previous living organism y Alternative is the theory of immutability of species o In favour of organic evolution o If 1 change can occur so can 2 or more changes JEAN BAPTISTE DE LAMARCK 1744-1829 y French naturalist; species are not unchangeable and that complex species evolved from simpler ones y DOCTRINE OF ACQUIRED CHARACHTERISTICS o Characteristics acquired are inheritable and can become part of legacy of succeeding generations; characteristics neglected tends to become vestigial CHARLES DARWIN 1809-1882 y THEORY OF NATURAL SELECTION o Competition between organisms and with environment results to preservation of favourable variations and rejection of injurious variations o Struggle for existence

JULIAN HUXLEY 1942 y MODERN SYNTHESIS Body of knowledge that now exists in evolutionary theory y Synthesis of basic premise y set forth by current genetic insight, natural selection and the findings of the different disciplines of life science PAEDOGENESIS y Phenomenon in larvaceans, agroup of protochordate sin which gonads become sexually mature and reproduction takes place in what is a larval organism y Back then gonads develop fast contrary to the maturation rate of other organs ; as a result larvaceans reproduce and die before they have fully metamorphosed NEOTENY y Analogous condition to paedogenesis found in Necturus tailed amphibian or urodele retains several larval gills and gill slits throughout life using them as respiratory organs y Perensnibranchiate species there is retardation of somatic feature, not precocious to development of gonads y Neotenic urodeles can be caused to resorb their gills and close their gill slits by use of thyroid hormone y Neoteny adults retain certain larval traits PAEDOMORPHOSIS y Phenomenon in which larval or immature features of ancestors become adult characteristics of descendants HETECHRONY y Paedogenesis, neoteny, paedomorphosis are manifestations of hetechrony y Change in relative rates of development of characters during phylogeny y One way of bringin about evolutionary changes y Basis fot the minority theory of phylogenetic origin of cephalochordates HISTORY OF ANATOMY y EGYPTIANS embalming y GALEN greek; dissected apes; 100 treatise on anatomy and medicine y LEONARDO DA VINCI y VESALIUS 1533 o De humani corporis fabrics on the structure of the human body  Hyoid and kidney from dog y PIERRE BELON dissected cetaceans & other marine animals vertebrate and invertebrate including fishes; aves y GEORGES CUVIER father of comparative anatomy and paleontology o Lecons d anatomie compare o Intro to CompaAna o Natural history of fishes y GEOFFROY SAINT-HILARIE y CHARLES DARWIN artificial and natural selection WORDS TO PONDER PRIMITIVE origin ; state of being ancestral GENERALIZED state of potential adaptability SPECIALIZED represents adaptive modification DERIVED/MODIFIED state of change from previous condition HIGHER & LOWER relative position of major taxa on phylogenetic scale SIMPLE no complex ADVANCED - modification because of further adaptation DEGENERATE- should be avoided VESTIGIAL phylogenetic remnant well developed in an ancestor

RUDIMENTARY - phylogenetic ; structures that became more exploited in descendants : ontogenetic ; structure that is not fully developed CHAPTER 3: PROTOCHORDATES AND ORIGIN OF VERTEBRATES PROTOCHORDATES Kingdom Animalia Phylum Chordata Subphylum Urochordata (Tunicata) Class Ascidiacea (sea squirts) Class Larvacea (Appendicularia) Class Thaliacea (Thaliaceans) Subphlyum Cephalochordata (Amphioxus) Branchiostoma or Asymmetron Subphylum Vertebrata (Craniata) UROCHORDATES y Notochord is confined to the tail y Known as TUNICATES, body is covered with nonliving tunic w/ cellulose like Tunicin, appears colored and transparent y Free swimming larval stage, only ASCIDIANS or sea squirts undergo complete metamorphosis , resorb notochord w/tail and become sessile adults y ALL are FILTER FEEDERS y THALIACEANS o Lack notochord and tail o Live far from shore among planktons o Cylindrical shape, linear colonies y LARVACEANS o Permanently larval marine tunicates o Body and long tail up to 8mm length o Entire organism surrounded by delicate gelatinous tunic bigger than the organism o Reaches sexual maturity during larval state paedogenesis y Larval Sea Squirts o 0.5 11.0 mm, fleeting existence lasts for only few minutes to few days o Do not feed o Body  Trunk w/ immature viscera  Muscular tail locomotion  Notochord stiff rod of small # of epithelial cells arranged i a single layer around central core of matrix  Filamentous sheath  36 - 1600 uninucleated striated muscle fibers unlike in vertebrates where they are multinucleated  No tendons  Sensory vesicle houses otolith and ocellus, brain  Otolith stimulates nerve endings for statoreception  Ocellus light sensitive pigment protected receptor cell  No definitive blood cells or heart till after metamorphosis  Branchial basket pharynx, where respiratory water enters via mouth over gills lining pharyngeal slits and enters the Atrium around the pharynx  Atriopore where oxygen depleted water is ejected  Adhesive papillae 3 appear at metamorphosis; w/ sticky secretions that attach larvae to permanent substrate y Adult Sea Squirts o At metamorphosis: Notochord is resorbed; nervous system is altered; viscera is rearranged o Incurrent siphon developed larval mouth o Excurrent siphon developed atriopore o Polyp is now a filter feeder

o Heart at each end there is now a functioning blood vessel ; blood is pumped alternately forward then backward; heart pauses before reversing the flow CEPHALOCHORDATES y Amphioxus sharp at both ends; lancelet y BRANCHIOSTOMA generic name y ASYMMETRON only other genus in the subphylum y Marine organisms; sand burrowers with only oral hood protruding for filter feeding y 2-8 cm in length, Branchiostoma californiense-South China y Living ones are semitransparent y Body o all trunk; metamerism of muscle segments or myomeres under thin skin > shaped under XS of body ; myoseptum separetes myomeres o Gill slits open to fluid-filled interior cavity Atrium o Atrium surrounds pharynx laterally and ventrally; collecting chamber for respiratory water,metabolic wastes and gametes; protects delicate gill filaments from abrasive sand action o Atriopore mid ventral exterior opening o # of pair of gill slits varies but exceeds 60 in adults o Tongue bar divide larval slit because of its downgrowth o Secondary gill bars result of division of larval slits o Skin is for respiration; single layer of epidermis and thin dermis; laraval skin ciliated for locomotion o Metapleural folds unknown function; longitudinal ridges o Notochord extends from the tip of rostrum to tip of tail; made up of muscular disks arranged in a long columns separated by fluid filled spaces o Dorsal hollow CNS hollow brain and dorsal nerve cord with central canal lined w/ Ependyma non nervous supportive membrane; caudal end is of ependymal cells alone o Leptomeninx connective tissue around cord and brain o Prosencephalon anterior cilia lined brain division o Deuteroencephalon posterior brain division o 39 or more pairs of branchial, oral and cranial nerves; 7 pairs of cranial nerves if branchial nerves is omitted o Spinal nerves merge from cord metamerically; made up of dorsal roots- sensory and motor fibers that supply only to atrial muscles and visceral organs & ventral roots not nerve roots, tubes that conduct extension of the body wall muscle cells into spinal cord where they receive innervations o Small brain correlated w/ scarcity of special organs: NO retinas, semicircular canals, lateral-line organs, olfactory epithelium o Chemoreceptors abundant on buccal cirri and velar tentacles, tail more sensitive than trunk o Tactile receptors elicit withdrawal present over entire body o Ocelli embedded on ventrolateral walls of spinal cord; consists of receptor cell and cap-like melanocyte o Vestibule collecting chamber for sea water o Oral hood laterally bounds vestibule o Velum - caudally bounds vestibule ;perpendicular membrane o Wheel organ stubby projections in the vestibule; covered with mucus to retrieve heavier food particles that miss the mouth o Buccal cirri partially strain water as it enters vestibule o Hypobranchial groove or endostyle -pharyngeal floor o Epibranchial groove pharyngeal roof o Peripharyngeal bands on gill bars; ciliated connect the 2 grooves o Midgut ring holds food particles

o Intestinal cecum evagination arised the same manner as human liver ; not homologous and differ in function; cecum secretes digestive enzymes for extra-cellular digestion w/in cecum and intestine and its lning cells phagocytose smallest food particles and digest them by intracellular digestion o Protonephridia excretory organs liealong 2ndary gill bars o Solenocytes - makes up protonephridium; project into coelom o Protonephridial chamber flagella of solenocytes project o Mature gonads visible through body wall and bulge into atrium, where sperm and eggs are shed larger gonads = ovaries; smaller gonads = testes; DIOECIOUS ovaries and testes do not develop in the same individual Arterial and Venous channels similar to fishes Amphioxus and vertebrates contrasted y No cephalization y No paired sense organs y w/ notochord but no vertebral column y w/ gill slits but in large numbers emptying into an atrium y dorsal hollw CNS but brain lacks divisions y segmented musculature but extend to anterior tip of head y median fins but no paired ones y 2 layered skin but epidermis is only 1 cell thick y Arterial and venous channels but no heart y Coelomate is restricted y Liquid wastes removed from coelomic fluid but excretory nephridia resembles non-chordates HEMICHORDATES y Incertae sedis of uncertain status y Acorn tongue worms marine organisms that live in the mud under shallow waters, fragile, 1.5 m length y Includes enteropneusts, class Pterobranchia and class Planctosphaeroidea y 1870 Karl Gegenbaur Enteropneusta for Acorn Worms y 1884 William Bateson added enteropneusts to phylum chordate o Dorsal strand of nerve cells and fibers w/in epidermis in a longitudinal groove on rthe dorsalsurface from collar region to end of the trunk. The groove sinks beneath the surface in the collar region and this becomes the collar nerve cord with lumen = manifestation of neural groove and neurocoel of embryonic chordates o Gill slits in lateral walls of endodermal foregut leading to exterior = homologous to pharyngeal slits o Short diverticulum of foregut, the stomochord extends forward into the proboscis = homologous to notochord of chordates o Echinoderms close kin if Enteropneusts  Bilateral symmetric larvae tornaria  Development process, muscle protein  Nervous system of enteropneust unlike dorsal hollow CNS of protochordates  Doubtful homology of stomochord and notochord  Enigmatic nature of enteropneust CNS ORIGIN OF VERTEBRATES y OSTRACODERMS o Strange little fishes 2- 30 cm long o No jaws,paired fin ; filter feeders o Broad bony plates in the skin form protective shield over head and much of the trunk hence ARMORED FISHES o Succeeded by jawed fishes o Bone in the skin of ostracoderms may have appeared w/ relative suddenness in some soft-bodied protochordate>Cephalochordates?

y Echinoderms are like vertebrates with mineralized skin in mesoderm and deuterostomous y Pikaia gracilens by C.D. Walcott o Soft bodied, laterally compressed, ribbin shaped fossils 5 cm long o resemble amphioxus zigzag myomeres, notochord that extends length of body y proposed prechordate o sessile or semisessile o bilaterally symmetrical o filter feeding o deuterostomous stem chordate o motile larval stage o dorsal hollow nerve cord o pharyngeal gill slits o notochord confined to larvae, where it is a stiff tail for locomotion o NEOTENY presence of notochord throughout life o EVOLUTIONARY CONVERGENCE explanation for similarities of protochordates and vertebrates AMMOCOETE: VERTEBRATE LARVA y Free swimming larvae of lampreys y Mistaken for adult protochordate y Lives from 2-6 yrs as a burrowing freshwater filter feeding larva with all basic chordate features and lacks vertebral column y Body o Dorsal hollow CNS  Brain w/ 3 vertebrate subdividion fore,mid,hind  Spinal chord o Notochord under hindbrain and extends to tip of tail o Pharynx 7 pairs of gill pouches opening exterior via gill slits o Endostyle complicated subpharyngeal gland ventral to pharynx & opens to short duct o Gland secretes mucus to trap food particles, degenerates when a larva metamorphoses since adult lampreys are not filter feeders o Body wall musculature arranged in myomeres that provide locomotion o Epidermis w/ many unicellular mucus glands o Dermis thin and fibrous o Simple digestive tract straight from mouth to anus o Stomodeum invagination of head ectoderm grows toward digestive tract develops into an opening then ultimately gives rise to buccal funnel and buccal cavity that opend to the pharynx o No stomach o Liver diverticulum w/ embedded gall bladder that disappears at metamorphosis when food changes from diatoms to tissue juices of prey o Kidneys located anteriorly to coelom made up of 3-6 tubules w/ funnell- like nephrostomes that collect coelomic fluid secreted by vascular tufts glomeruli o Rudimentary gonad paired in early larvae but unpaired in later ones o Circulatory system is typically vertebrate  Heart - sinus venosus, atrium and ventricle o Special sense organs  Median olfactory sac receptor for smell  2 middorsal eyes pineal and parapineal organs made up of simple lens and receptor cells  Paired lateral eyes lie deep under skin, vestigial  Otic vesicles invagination of head ectoderm and develop to membranous labyrinths of the inner ears

 Neuromast organ lie on skin of head & open to surface via pores ALTERNATE THEORY OF AMMOCOETE-PROTOCHORDATE RELATION HETEROCHRONY y An isolated population of ammocoete-like larvae, a genetic differential in the rate of development of the larval gonads had become established such that gonads matured before metamorphosis and reproduction of larvae become possible, results to a protochordate similar to amphioxus y Cephalochordates are from agnathan like ancestor the larvae through heterochrony became sexually mature giving rise to a new species ; explains the similarities between amphioxus and ammocoete larvaa y Resemblance may be explained through parallel or convergence evolution y Objections include lack of agnathan or other vertebrate larvae of an atrial chamber alongside the gills CHAPTER 4: PARADE OF VERTEBRATES IN TIME AND TAXA VETEBRATE TAXA y Conventional natural classification Mammalia Amniotes Aves w/ extraembryonic Tetrapods membrane amnion Reptilia Amphibian Gnathostomes w/ jaws Ostechthyes Chondrichthyes Anamniotes Placodermi Lack amnion Fishes Acanthodii Agnasthomes Agnatha
Lack jaws

AGNATHA jawless fishes y OSTRACODERMS ancient bony ; late Cambrian o Entire body covered by bony dermal armor made up of broad plates and smaller tile-like scales o No jaws and paired fins although some had STABILIZERS bizarre projections from body wall o Lived largely on fresh or brackish water by late Silurian some were marine o E.A. Stenso palaeontologist o Head skeleton more or less flat spicule covered bony shield w/ 4 dorsal apertures; 2 dorsal apertures upward-staring eyes, 1 median or pineal eye and single naris where nasopophyseal duct led to an olfactory sac o Heterocercal (unequally divided tail) like most paleozoic fishes o Disappeared at end of Devonian y CYCLOSTOMES extant eel like boneless lampreys and hagfishes o Highly specialized  Lost bone everywhere in the body  Adaptations for parasitism buccal funnel and rasping toungue  Absence of recognizable vertebrae  No centra around notochord, chief skeleton of trunk and tail o Primitive features  Median nostril connected with a single olfactory sac  No paired fins and skeletal elements  2 pairs of semicircular ducts instead of three found in higher vertebrates o PETROMYZONTIFORMES - Lampreys  Ammocoete larvae  Large buccal funnel lined w/ horny denticles

Tongue like cartilaginous rod covered with horny teeth 1 dorsal nostril and a nasophypophyseal sac 7 pair of gill pouches Anadromous lampreys Petromyzon marinus marinus/dorsatus  Lampetra freshwater lampreys o MYXINIFORMES hagfishes  slime eels, marine cyclostomes w/ shallow buccal funnel lacking denticles  bottom feeding scavengers diet  parasitize slow moving fishes like lampreys  Myxine glutinosa atlantic hafish 6 pairs of gill pouches  Bdellostoma stouti 10-15 pairs of gill pouches  Not anadromous ACANTHODII y Oldest jawed fishes like ostracoderms head and body protected by dermal armor of bony plates and scales y spiny sharks skeleton made up of bone and cartilage y Large operculum, paired fins w/ 5 accessory pairs supported by hollow spines PLACODERMI y Armoured fishes abundant in freshwater; paired fins y Swift predators y Arthodires Coccosteus heavy bony dermal shield all over head and gill region, trunk then other pars covered by bony scale or left naked y Antiarchs - Bothriolepis small w/ atypical pectoral fins, dorsal eyes, flat ventral surface suggests bottom feeders CHONDRICHTHYES cartilaginous fishes y No bone in their body other than scales and teeth; mouth on ventral surface rather than terminal except in paleozoic sharks Cladoselache 7 gill slits y Placoid scales made up of basal plate and bony spine of dentin y Pelvic fin of males modified to form claspers used in reproduction y Internal fertilization; macrolecithal eggs y Oviparous species eggs encased in horny or leathery shell w/ tendrils that entwines around vegetation y ELASMOBRANCHS sharks, rays, skates and saw-fishes o Order Cladoselachii paleozoic sharks extinct o Order Pleuracanthodii paleozoic sharks extinct o Order Squiliformes modern sharks o Prder Rajiformes - modern skates, rays and saw fishes o 5 ventral naked gill slits Hexus - 6 Heptanchus 7 st o w/ spiracle enterior to 1 gill slit w/ pseudobranch or gill like surface o heterocercal caudal fin o Squalus acanthias viviparous atlantic spiny dogfish w/ prominent horny spine o Squalus suckleyi - pacific spiny dogfish o Mustelus smooth dogfish shark lacks dorsal spine o Dorsoventrally compressed w/ anterior fins attached to head and trunk forming wing like organs y HOLOCEPHALANS chimaeras or ratfishes o Chimaeras lack scales; fleshy operculum hides gill slits; closed spiracle o Distinct upper jaw solidly fused w/ cartilaginous braincase o Have hard flat bony plates on jaws

   

OSTEICHTHYES - bony fishes y Bony skeleton; gill slits covered by bony operculum; terminal mouth; gas-filled swim bladder; shallow adult cloaca virtually nonexistent y ACTINOPTERYGII RAY FINNED FISHES o Slender fin rays are sole support for otherwises membranous fins; lack internal nares o Bony armor covered w/ enameloid called ganoin; heterocercal caudal fin o Superorders Chondrostei paleozoic/ paleoniscoids  Order Acipeseriformes y Sturgeons, spoonbills or paddle fishes o Cartilaginous endoskeleton; small bony scale on tail  Order Polypteriformes y Polypterus and Calamochthyes - African freshwater lungfishes o Ganoid scales, ossified endoskeleton o Superorder Holestei freshwater fishes  Lepisosteus - garfishes; lepisosteoid ganoid scales  Amia calve bowfin; modern head covered / bony platelike scales lacks ganoin; trunk and tail have modern fish scales o Superorder Teleostei  Tail no longer heterocercal; scale s no longer heavily bony and inflexible  Thinner dermal bones; movable jaws and palate  Pelvic fins far forward  95% of all living fishes y SARCOPTERYGI - LOBE-FINNED FISHES o Prominent fleshy lobe at base of paired fins; lobe w/ fin skeleton o w/ internal nare open to oropharyngeal cavity o Order Crossopterygii ancestors of amphibians  Common Devonian now extict Latimeria  Skeletal elements; early tetrapod limbs st  Skull similar to 1 amphibians  Sub order Rhipidistia freshwater  Suborder Coelacanthini Latimeria ; lacks internal nares o Order Dipnoi true lungfishes  Inefficient gills  Protopterus Africa  Lepidosiren Brazil  Neoceratodu Australia AMPHIBIA y LABYRINTHODONTS oldest tetrapods o Swamp-dwelling; named because of dentin of their teeth was infolded to resemble a labyrinth when viewed XS o Minute bony scales in dermis; fishlike tail w/ dermal fin rays o Operculum and sensory canal system of neuromast organs like fishes o Order Ichthyostegalia oldest devonian o Order Temnospondyli Permian o Order Anthracosauria paleozoic direct ancestors of reptiles y LEPOSPONDYLS upper carboniferous amphibians o Salamander like amphibians; extinct o Orders lacked limbs, some resemble urodeles and others w/ bizarre triangular skulls o Skull study shows that they may have come from crossopterygian ancestor independent from labyrinthodonts and urodeles ma be descendants of specific lespondyls y LISSAMPHIBIANS Triaissic and modern amphibians o Order Anura frogs, toads and tree toads  Tail-less w/ caudal vertebrae fused into one elongated urostyle; breathe w/ lungs and skin

 Jelly encapsuled eggs in ponds; larval stage lacks aquatic environment  Earlies known anuran Triasobatrachus = Protobatrachus Triassic  Family Ranidae long legged slender bodied frogs  Family Bufonidae terrestrial toads w/ parotid glands  Family Hylidae tree frogs o Order Urodela tailed amphibians  Perennibranchiate  Necturus, Proteus, Ambystoma, Siren, amphiuma , Hynobius, Cryptobranchus, Salamandra, Notopthalmus, Plethodon, Pseudobranchus o Order Apoda - legless burrow-dwelling species  Circumtropical limbless except for aquatic species living in burrows in swampy places REPTILIA y Cotylosaurs - stem reptile y Sphenodon lizard like relict y Amnion, chorion and allantois extraembryonic membranes y Amnion fluid filled membranous sac y Chorion and allantoids vascularised membrane that lies against porous egg shell; replaces larval gills for respiration y Oviparous reptiles; cornifies epidermal cells plaques shields or surface scales y Neck, occipital condyle, pelvic girdle and 2 sacral vertebrae, claws, ,metanephrous kidney y Hert divided to left and right chambers separating systemic and pulmonary circulation y Ectoherms- cannot maintain more or less constant body temperature y Scaly, clawed, mostly terrestrial tetrapods lacking feather and hair except viviparous species lay macrolecithal shell covered cledoic eggs on land, embryos develop w/in amnion and young hatches fully formed y Subclass ANAPSIDS cotylosaurs and turtles o No bony temporal arch; bony dermal plates, loss of trunk musculature; no teeth o Order Testudinata turtles o Order Chelonia turtles y Subclass LEPIDOSAURS o Order Rhynchocephalia Sphenodon  Tuataras primitive lizard like reptile but w/ diff. teeth, scale and internal morphology o Order Squamata lizards, snakes and amphisbaenians; have only remnants of one or both arches; loss is correlated w/ expansion of jaw muscles  Suborder Lacertilia lizards y Most versatile; well developed appendicular muscles y Spectacles transparent eyelids rd y Nictitating membrane -3 eyelid  Suborder Amphibaenians snakelike burrowers y Subterranean lizards, limbless w/ annulated bodies y Eardrum and eyes covered w/ opaque skin  Suborder Serpentes snakes y Evolved from lizards, limbless, viviparous o Primitive have 2 temporal fossae hence DIAPSID y Subclass ARCHOSAURS Mesozoic vertebrates ; Diapsid skulls o Subclass Thecodontia stem archosaurs; teeth set in deep sockets o Subclass Pterosauria flying reptiles; w/ pneumatic bones th and batlike wings supported by elongated 4 finger, long neck and tail o Subclass Saurischia dinosaurs w/ reptilian pelvis; swift predatory carnivores

o Subclass Ornithiscia dinosaurs w/ birdlike pelvis; herbivores o Subclass Crocodilia sole surviving archosaur; slender th triangular snout and enlarged 4 tooth y Subclass EURYAPSIDS - extinct marine reptiles o Parapsida and Synaptosauria o Ichthyosaurs fislike w/ no visible necke o Plesiosaurs long neck, short flat trunk, short tail, paddlelike limbs o Accommodate reptiles w/ single dorsal temporal fossa y Subclass SYNAPSIDS o Reptiles from which mammal emerged; distinct single lateral temporal foss o Pelycosaurs earliest synapsid; parietal foramen w/ median eye o Therapsida - mammal like AVES y Endothermic vertebrates w/ feathers from bipedal ornithiscian dinosaur y Endothermy ability to maintain stable body temp. Despite environment or warmblooded y Carina large sterna keel for attachment of massive flight muscles absent in most birds that do not fly y ARCHAEORNITHES oldest birds known Archaeopteryx y NEORNITHES o Odontognathae w/ teeth  Hesperonis vestigial wings but swam for fish; sharp conical teeth o Neognathae toothless  Ratites cannot fly; powerful leg muscles y Ostriches, emus, cassowaries, rheas, kiwis, penguins  Carinates can fly y Annual migrants MAMMALIA y Amniotes w/ synapsid skull, hair except in monotremes, mammary glands and nipples; single dentary bone on each side of lower jaw; 3 bones in middle ear cavity, muscular diaphragm separating thoracic and abdominal cavities; sweat glands; 2 sets th of teeth; loss of 4 aortic arch; sound collecting lobe; specialized larynx; development of cerebral cortex y Subclass Protheria lay eggs and have cloaca throughout life o Order MONOTREMATA  Ornithorhyunchus platypus or duckbilla  Anteaters or echidnas long sticky tongues & stout claws; armed w/ sharp quills  No nipples but milk exude from modified sweat glands y Subclass Metatheria viviparous mammals w/ yolk sac as placenta o Order MARSUPIALIA  Fetal yolk sac in contact w/ chorion serves as placenta  Young born in larval state then nursed in marsupium or maternal abdominal pouch  Kangaroos, wallabys, Tasmanian wolves dog like, bandicoots- rabbitlike rats, wombats, phalangers, moles  Resemblance of phenotype w/ placental mammals are instances of parallel evolution y INSECTIVORA flying lemur, mole, solenodon, hedgehog, shrew o Flat footed, 5 toes, smooth cerebral hemispheres, small sharp pointed teeth, large embryonic allantois, shallow cloaca, testes in abdominal cavity y CHIROPTERA bats o From primitive insectivore

o Paragium wing; double membrane of skin stretched along length of body bet. Trunk, forelimb and hindlimbs extending to the tail o strong pectoral muscles and keeled sternum o Insectivorous, frugivorous or sanguinivorous PRIMATES o Specializations grasping hand, nails, opposable big toe, prehensile tail, cerebral hemispheres, short snout, one pair of nipples o Primitive flat footed gait, 5 digits, large clavicle, central wrist carpal, generalized dentition o Suborder Lemuroidea lemurs and lorises  Lemurs - Long tail, 2nd finger and toe have claw instead of nails, duplex uterus, nondeciduate placenta  Lorises slow moving nocturnal primates. No tail and index finger is vestigial; related to pottos and bush babies o Suborder Tarsioidea tarsiers and relatives  Tarsiers resemble higher primates, eyes are close together and directed forward, head is balanced at right angles to vertebral column, all 5 fingers have nails and so do toes except 2nd and 3rd, deciduate placenta o HIGHER PRIMATES  Suborder Platyrrhini monkeys and marmosets y Noses open to the side y Cebus - capuchin monkeys y Ateles spider monkey y Alouatta howler monkey; enlarged hyoid bone and larynx  Suborder Catarhini old world monkey, apes, man y Anthropoids ; nostrils close together & open downward y Superfamily Cercopithicoidea baboons,mandrils, macaque or rhesus monkey y Superfamily Hominoidea chimpanzees, gibbons, gorillas, orang-utans and man ALL have no tail  Skull is set at right angle to vertebrae; 32 teeth, deciduate placenta CARNIVORA flesheaters o Powerful jaws, long sharp canine teeth, highly convoluted cerebral cortex, capable of considerable learning, most have 5 toes , sharp retractable claws o Suborder Fissipedia terrestrial  Cats,civets, hyenas, dog-like carnivores, bears and giant pandas, racoons, mustelids o Suborder Pinnipedia marine  Family Phocidae true seals(earless wriggling seals)  Family Otaridae eared seals (fur and sea lions)  Walruses CETACEA whales, dolphins, porpoises o horizontal tail w/ 2 lobes or flukes composed of fibrous connective tissue, paddle like forelimbs for balance, vestige hind limbs and girdles , single large blowhole, blubber, good eyesight, scan environment through echolation, great sense of taste, no sense of smell EDENTATA insectivorous mammals o Sloths and anteaters, armoured edentates Armadilloidentical quadruplets from single zygote o No incisor or canine and cheek teeth, some front clas TUBULIDENTATA - Aardvarks o Another order of anteaters, elongated piglike snout; long strong claws; long sticky tongue o Teeth is a major criteria in mammalian taxonomy o Teeth are peg-like cylinders w/ shallow roots PHOLIDOTA - Pangolind o Toothless scaly anteaters Manis

y RODENTIA o Largest mammalian order o Single pair of long curved incisor teeth on each jaw; enamel on outer surface of teeth; no canine teeth so with diastema; cellulose eaters o Suborder Sciuromorpha squirrels, chimpunks, woodchucks, gophers, beaver o Suborder Myomorpha micelike rodents, rats, voles, hamsters, lemming o Suborder Caviomorpha/Hysricomorpha porcupines, cavys, nutria / coypus, chinchillas y LAGOMORPHA o Pikas amd Hare & Rabbits o All are herbivores; 2 pair of incisors on upper jaw o Rabbits differ from hares in being born in fur-lined nest, blind, virtually hairless and helpless o Hares are born w/ eyes open and able to run after few minutes o Pikas have smallerbody, shorter ears and equal length of limbs y UNGULATES walk on tips of their toe sprotected by hoofs; no clavicle; w/ horns; herbivores o Order PERISSODACTYLA  horses and horselike mammals, tapirs and rhinoceros  Most of body weight is borne on a single digit mesaxonic foot  Odd-toed ungulates, but tapirs and rhinos have 4 toes o Order ARTIODACTYLA  Body weight borne by 2 toes paraxonic foot  Even number of toes  Pigs, hippopotamuses, peccaries, cattle, camels, llamas, deer, antelope, giraffes y SUBUNGULATES - herbivores o Order PROBOSCIDEA  Elephants, mastodons  Proboscis, scanty hair, thick wrinkled skin; tusks, no canin teeth, large molars; bulky animals, 5 toes w/ thick hooflike nails, elastic pad on back of each toe bears body weight o Order HYRACOIDEA  Hyraxes; short ears; hunched body at rest; harelip; incisor teeth grows continuously; plantigrade or flatfooted; 4 fingers 3 toes; ungulate molar teeth o Order SIRENIA  Sea cows; stout vegetarian manatees and dugongs

VARIATION AMONG INDIVIDUALS y Homozygous genotype types possible w n pair is 2n y Number of diff. kinds of genotype possible is 3n y Genetic variation coupled w/ geographical isolation is the matrix for speciation to occur and for evolution to flourish

CHAPTER 5: EARLY VERTEBRATE MORPHOGENESIS Egg type Accdg. to amount of yolk Microlecithal very little Mesolecithal moderate Macrolecithat massive Oviparo Accdg, distribution w/in egg Isolecithal even distribution Telolecithal yolk mass tends to concentrate on the vegetal pole while the animal pole is yolk free