- PHYLUM ECHINODERMATA - . GENERAL CHARACTERISTICS
Transcription
- PHYLUM ECHINODERMATA - . GENERAL CHARACTERISTICS
- . Around t h e i r base a r e many modified w i t h an epidermis. spines, the p e d i c e l l a r i a , which resemble t i n y jaws o r scissor blades mounted on a s t a l k . When stimulated mechanically o r chemically, the p e d i c e l l a r i a may be opened by muscles. Their f u n c t i o n i s t o keep t h e aboral surface clean, t o a i d i n t h e capture o f small food p a r t i c l e s and p r o t e c t the dermal papulae which a r e t h i n walled extensions o f the coelom and f a c i l i t a t e gaseous exchange. PHYLUM ECHINODERMATA -. - " GENERAL CHARACTERISTICS Spiny-skinned animals possessing calcareous o s s i c l e s embedded i n t h e dermis and spines p r o t r u d i n g from the surface: t h i s creates an i n t e r n a l skeleton. They are r a d i a l l y symnetrical i n the a d u l t phase and a r e characterized by having f i v e rays, o r m u l t i p l e s o f f i v e (pentamerous). This penta-symnetry i s most apparent i n s t a r f i s h and Least apparent i n sea cucunbers. WATER VASCULAR SYSTEM The water vascular system i s a d i v i s i o n o f t h e coelom and i s unique t o echinoderms. I t i s e s s e n t i a l l y a system o f hollow interconnected tubes f i l l e d w i t h water. The water i s under a s l i g h t p o s i t i v e pressure. Water enters the system v i a the madreporite on t h e aboral disc. The water vascular system terminates i n a t u b e - f e e t (Podia) i n the grooves on the o r a l surface. The sequence o f canals i n t h e water vascular system i s as follows: Echinoderms have n o t always been r a d i a l l y symnetrical: f o s s i l forms a r e b i l a t e r a l l y symnetrical ( w i t h a l e f t and r i g h t sides and a f r o n t and back). Echinoderm Larval stages ( b i p i n n a r i a ) a r e a l s o b i l a t e r a l l y symnetrical. Body w a l l s a r e cornposed o f t h r e e layers o f c e l l s . Most tissues a r e c i l i a t e d . There i s no head region, b r a i n nor segmentation. 1 Madreporite (sieves the water) I Stone Canal (runs downwards) 1 Ring Canal ( e n c i r c l e s the mouth) 1 5 Radial Canals (one per r a y arm) 1 Numerous Transverse Canals (perpendicular t o r a d i a l cana 1) 1 Ampullae: The head o f the tube f e e t . A l l examples a r e marine and t h e r e a r e no p a r a s i t i c forms. ALL have tube f e e t which extend by h y d r a u l i c pressure from t h e animal's water vascular system, which are used v a r i o u s l y f o r Locomotion, burrowing, exchange o f gases ( r e s p i r a t i o n ) and feeding. The water vascular system i s a h y d r a u l i c pressure system. The s t a r f i s h moves by means o f i t s tube feet. A tube f e e t elongates as the muscles surrounding the ampullae contract f o r c i n g f l u i d i n t o the foot. On t h e bottom o f the f o o t i s a sucker which adheres t o t h e substratum ( s t i c k secretions from the gland c e l l s a t t h e t i p a i d i n adhesion). Longitudinal muscles then contract, shortening the r a y arm and h e l p p u l l t h e s t a r f i s h forward. The combined e f f e c t s o f forward t h r u s t s o f many co-ordinated tube-feet produce Locomotion i n the starfish. The d i g e s t i v e t r a c t i s simple u s u a l l y complete (but may Lack an anus and t h e r e f o r e be incomplete). Respiration i s achieved by minute g i l l s (dermal papulae) which protrude from t h e coelom, by tube f e e t and i n the case o f sea cucumbers by a cloaca1 r e s p i r a t o r y trees. A c i r c u l a t o r y system r a d i a t e s from t h e c e n t r a l coelom, the coelom i s Lined w i t h c i l i a t e d peritoneum and i s u s u a l l y Large and i t s f l u i d contains f r e e amoebocytes. Part o f t h e Larval coelom becomes a water vascular system u s u a l l y having many tube feet, serving f o r Locomotion, food handling, o r r e s p i r a t i o n . Tube f e e t are a l s o used f o r c a p t u r i n g and handling food, and f o r r e s p i r a t i o n and excretion. There a r e a l s o n i n e small spherical swellings c a l l e d t h e tiedemannls bodies on the inner r i n g canal. 2 The nervous system c o n s i s t s o f a circum o r a l r i n g and r a d i a l nerves i n t o each arm. The sexes a r e separate ( r a r e exceptions), and a l i k e externally; gonads a r e Large w i t h simple ducts; ova are abundant. The l a r v a i s free-swimning, and i s characterized by a conspicuous metamorphosis t o become adults. A few species are viviparous, a few reproduce asexually by s e l f - d i v i s i o n , and many regenerate Lost parts readily. For example s t a r f i s h are generally capable o f r e p l a c i n g one o r more Lost arms; sometimes, even a s i n g l e arm can regenerate i n t o an e n t i r e animal. GENERAL STRUCTURE OF A STARFISH (ASTEROIDEA) EXTERNAL The Lower surface w i t h t h e mouth i s c a l l e d the o r a l surface. The upper surface w i t h the anus i s c a l l e d the aboral surface. Both t h e mouth and anus are c e n t r a l l y situated. ORAL SURFACE Has f i v e ambulacral grooves, one i n each r a y from which extends two o r four rows o f tube-feet. The r a y may be f l e x e d s l o w l y by a few r e t r a c t o r muscles i n the body wall. ABORAL SURFACE Many spines o f various sizes, p e d i c e l l a r i a e and dermal papulae a t t h e base o f t h e spines, a madreporite which i s the entrance t o the water vascular system and an anal opening. DIGESTIVE SYSTEM + This i s a short b u t complete system I Mouth 1 Short oesophagus 1 Thin w a l l stomach l a r g e Cardiac stomach small aboral P y l o r i c I From the P y l o r i c a tube passes i n t o each r a y 1 I n each r a y arm i t d i v i d e s i n t o two branches hepatic cecae from here I many Lateral pouches e x i s t . I Above the stomach i s a slender i n t e s t i n e 1 s p l i t s i n t o two branched pouches, the r e c t a l i n t e s t i n a l cecae 1 Anus -r -r stomach arm. called and FOOD S t a r f i s h eat almost any animal matter, and are a l s o carnivorous on oysters, mussels, barnacles, clams etc. SMALL FOOD passes d i r e c t l y p e d i c e l l a r i a o r tube feet. to the mouth via the BIVALVES the s t a r f i s h opens the s h e l l halves s l i g h t l y using i t s rayiarms and tube f e e t . The cardiac stomach i s everted between t h e gap i n t h e s h e l l halves. D i g e s t i v e enzymes are released and t h e prey t i s s u e i s broken down i n t o a f l u i d which i s swept i n t o t h e h e p a t i c cecae by the action o f c i l i a . N u t r i e n t s a r e s t o r e d i n t h e hepatic ceca. INTERNAL SYSTEM INTERNAL TRANSPORT An endoskeleton o f calcareous p l a t e s o r o s s i c l e s i s bound together by muscles and connective tissue. Uses the f l u i d mediun o f t h e p e r i v i s c e r a l coelom. The aboral calcareous spines are short, b l u n t and covered Amoebocytes (coelomocytes) a s s i s t i n the transport o f 5 n u t r i e n t s and wastes. rows o f tube-feet they are d i s t r i b u t e d b i l a t e r a l l y . C RESPIRATION SEA LILIES CLASS CRINOIDEA: occur as f r e e l i v i n g forms (feather s t a r ) o r s e s s i l e forms (sea L i l i e s ) . These animals are h i g h l y c a l c i f i e d , and generally occur i n deep waters. They c o l l e c t food by c i l i a r y currents set up on t h e i r complex arms. Tube f e e t are present only as small p a p i l l a e on t h e sides o f the ambulacral grooves. -- Gaseous exchange between dermal papulae and tube feet. Dermal papulae are outpockets o f t h e coelom t h a t pass through minute openings i n t h e skeleton. They are covered w i t h c i l i a on the i n t e r n a l and external surfaces. The external c i l i a keep a current o f oxygenated water passing over the outside, and t h e i n t e r n a l c i l i a c i r c u l a t e t h e coelomic f l u i d i n t o t h e papulae. SEA DAISIES CLASS CONCENTRICYCLOIDEA: c o n s i s t o f a d i s c The water shaped body without arms, mouth o r anus. vascular system possesses two r i n g canals concentric on the v e n t r a l surface. EXCRETION = - e Amnonia i s t h e p r i n c i p a l s o l u b l e nitrogenous waste, and i t d i f f u s e s through t h i n areas o f t h e body surface, e s p e c i a l l y tube f e e t and dermal papulae. Particulate waste are picked up by the amoebocytes w i t h i n the coelom; these amoebocytes are subsequently eliminated through the walls o f t h e dermal papulae. NERVOUS SYSTEM AND SENSE ORGANS This consists o f interconnected peripheral and c e n t r a l components. The p e r i p h e r a l nervous system i s composed o f a subepidermal nerve net t h a t extends throughout the body, i n c l u d i n g t h e tube feet. This system c o n t r o l s l o c a l responses o f t h e pedicel lariae, spines and dermal branchiae t o s p e c i f i c s t i m u l i . L a t e r a l t r a c t s conduct neural impulses a t higher speeds t o and from the c e n t r a l nervous system. This consists o f a nerve r i n g around the mouth plus f i v e l a r g e r a d i a l nerves t h a t run p a r a l l e l t o t h e r a d i a l canals o f t h e water vascular system. c 2 - Most echinoderms respond t o touch, gravity, l i g h t and chemical s t i m u l i . Yet t h e sense organs are poorly developed. The primary receptors are sensory c e l l s Located i n t h e epidermis, e s p e c i a l l y on t h e podia and t h e margins o f t h e ambulacral grooves. At t h e end o f each ray i s a small t a c t i l e t e n t a c l e and a l i g h t s e n s i t i v e eyespot composed o f 80 t o 200 o c e l l i . Most echinoderms are negatively phototaxic and generally seek shade. REPRODUCTION AND REGENERATION Sexes are separate. A p a i r o f branched gonads L i e i n the p e r i v i s c e r a l coelom a t t h e base o f each arm. Female s t a r f i s h may release up t o 2,5 m i l l i o n eggs a t one time. Males produce even more sperm. F e r t i l i z a t i o n and development occurs external l y i n t h e seawater. Echinoderm development: f e r t i l i z e d c e l l * cleavage = two-egg stage -r b l a s t u l a * gastrula * b i p i n n a r i a * branchiolaria * undergoes metamorphosis * pentasymnetrical s t a r f i s h . OTHER ECHINODERMS BRITTLE STARS CLASS OPHIUROIDEA: possess f i v e arms r a d i a t i n g from a c e n t r a l disc, the arms are more sharply formed. Tube f e e t Lack suckers, movement effected c h i e f l y by movement o f t h e arms brought about by muscles which connect successive ambulacral ossicles. They are d e t r i t u s feeders, having a capacious stomach but Lack any extension o f t h e gut i n t o t h e arms and Lack an anus. SEA URCHINS CLASS ECHINOIDEA: have a r i g i d body wall o r t e s t composed o f c l o s e l y f i t t i n g calcareous p l a t e s embedded i n the dermis. The o r a l surface i s morphologically very Large compared t o t h e aboral surface. The mouth i s surrounded by powerful jaws which can scrape o f f encrusting organisms from t h e rocks. The spines borne on t h e t e s t are Long and movable and are interspersed w i t h p e d i c e l l a r i a . The numerous tube f e e t are very e x t e n s i b l e and p r o j e c t beyond the spines. SEA CUCUMBERS CLASS HOLOTHUROIDEA: possess reduced dermal ossicles. They feed on d e t r i t u s , which they take i n t o the mouth by Large modified tube f e e t surrounding i t . The remaining tube f e e t resemble those o f the s t a r f i s h and are used f o r Locomotion. Although there are f i v e NOTE: CLASSES ASTEROIDEA and OPHIUROIDEA are sometimes considered t o be subclasses i n the CLASS STELLEROIDEA PHYLUM CHAETOGNATHA ARROW WORMS Arrow worms, are named f o r t h e i r streamlined appearance. They are about 40 mn i n Length and completely transparent b i l a t e r a l l y symnetrical deuterostomes. Their r e l a t i o n s h i p t o echinoderms, hemichordates o r chordates i s unclear. Arrow worms are important planktonic predators, feeding voraciously on copepods, f i s h f r y and each other; they capture t h e i r prey using b r i s t l e s ( c h a i t e ) around t h e i r They possess a hood which they can draw jaws (gnathos). over t h e i r b r i s t l e s and jaws t o streamline t h e i r bodies. Swimning i s due t o the a l t e r n a t i n g contractions o f the l o n g i t u d i n a l muscles on each side. The f i n s are f i r m l y attached t o t h e i r Lateral sides o f t h e body. The digestive t r a c t i s straight. The nervous system comprises a v e n t r a l ganglia i n t h e trunk and a dorsal ganglia i n the head region and several smaller ganglia. Sensory b r i s t l e s d i s t r i b u t e d over t h e body detect prey, and a c i l i a r y Loop t h a t passes beneath t h e head Like a necklace i s thought t o detect water currents o r chemicals. Two dorsal eyes ( o c e l l i ) presumably detect Light and c o n t r o l the v e r t i c a l m i g r a t i o n a t dawn and dusk. Arrow worms are hermaphroditic, w i t h two ovaries i n the trunk coelom and two testes i n t h e t a i l coelom. Sperm are packaged i n t o spermatophytes and are released by simply breaking through t h e body wall. These spermatophores then attach t o the a n t e r i o r trunk o f the same o r a d i f f e r e n t individual, and t h e sperm migrate i n t o t h e ovaries. F e r t i l i z a t i o n i s therefore internal. There are no excretory, c i r c u l a t o r y o r r e s p i r a t o r y organs. PHYLUM HEMICHORDATA This group o f animals share several s i m i l a r i t i e s w i t h the chordates which includes pharyngeal s l i t s and a dorsal nerve cord and were once c l a s s i f i e d as chordates since i t was thought t h a t they a l s o possessed a notochord. Recent analysis has i d e n t i f i e d what was once considered a notochord t o be an extension o f the buccal c a v i t y ( t h e buccal d i v e r t i c u l u n ) . Nevertheless t h e hemichordates appear t o have diverged close t o a p o i n t a t which the chordates originated. Hemichordates are d i v i d e d i n t o two classes t h a t d i f f e r g r e a t l y from each other, although they show s i m i l a r s t r u c t u r e i n e a r l y embryological stages. Class Enteropneusta includes most o f the phylun, and are comnonly r e f e r r e d t o as acornworms. These animals burrow i n marine sediments and have worm-like bodies d i v i d e d i n t o a proboscis, c o l l a r , and trunk. The other Class Pterobranchia Live i n colonies and secrete tubes. ACORN WORMS GENERAL FEATURES Possess a prominent proboscis, a c o l l a r and a long slimy trunk up t o 2 metres i n Length. Each o f these three regions have a separate coelomic compartment f i l l e d with spongy t i s s u e that provides some mechanical support, nevertheless these animals are e a s i l y broken. Acorn worms e i t h e r Live i n U-shaped burrows i n shallow water o r they burrow through marine sediment o r Live under rocks o r seaweed. . A ,'.The proboscis i s the main organ o f Locomotion, with the trunk being p u l l e d along passively. Many acorn worms feed by i n g e s t i n g Large q u a n t i t i e s o f mud o r sand from which the gut e x t r a c t s organic debris. Others feed by means o f c i l i a on t h e proboscis which pass the food backwards i n t o the mouth. Food p a r t i c l e s are bound on a mrcous s t r i n g and are swallowed along w i t h the water. The chordates are b i l a t e r a l l y symnetrical, w i t h three germ Layers, a segmented body, complete d i g e s t i v e t r a c t , and w e l l developed coelom. The f o l l o w i n g three c h a r a c t e r i s t i c s apply:- * A s i n g l e dorsal t u b u l a r nerve cord. A notochord. ACORN WORM INTERNAL STRUCTURE G i l l s l i t s i n the pharynx Uater swal lowed d u r i n g feeding e x i t s through pharyngeal s l i t s and g i l l pores. The pharyngeal s l i t s are U-shaped openings i n the pharynx t h a t can swallow water i n t o the pharyngeal pouches and out through the g i l l pores. Pharyngeal s l i t s are sometimes c a l l e d g i l l s l i t s , although i n these animals they have a feeding r o l e rather than f o r t h e exchange o f r e s p i r a t o r y gases. The body : surface i s t h e main route f o r r e s p i r a t o r y exchange. The colourless blood i s punped a n t e r i o r l y by a dorsal vessel i n t o a c o n t r a c t i n g heart vessel i n the proboscis and then p o s t e r i o r l y by a v e n t r a l vessel. The blood c i r c u l a t e s ; through sinuses, making i t an open c i r c u l a t o r y system. One such sinus, the glomerulus, i s assuned t o have an excretory function. The nervous system consists Largely of a d i f f u s e network i n the base o f the epidermis. Along the dorsal and v e n t r a l midlines t h i s plexus i s concentrated i n t o dorsal and v e n t r a l nerve cords which Lack ganglia. I n places the nerve cord i s hollow and s i m i l a r t o the hollow d o r s a l nerve o f the chordates. Sensory receptors are s c a t t e r e d over the integunent, e s p e c i a l l y around the proboscis. Sexes are separate i n , the acorn worms, although t h e r e i s L i t t l e t o d i s t i n g u i s h males from females. The gonads have a separate gonadopore, and f e r t i l i z a t i o n i s external and achieved through mass spawning i n i t i a t e d by the females. Embryological development i s t y p i c a l deuterostome. This i s characterized by r a d i a l cleavage, the blastopore becoming the anus, and the coelom forms and outpocketing o f the archenteron. Development i s sometimes d i r e c t , although more u s u a l l y a t o r n a r i a Larva develops characterized by c i l i a r y contractions which cause i t t o spin. The t o r n a r i a Larva c l o s e l y resembles the b i p i n n a r i a n Larva o f the echinoderms. PTEROBRANCHIA There are about t e n species i n t h i s class and they have a s i m i l a r development t o the acorn worms w i t h t h e i r bodies d i v i d e d i n t o a proboscis, c o l l a r and trunk. However, the adult, external appearance i s very d i f f e r e n t from acorn worms. The c o l l a r expands d o r s a l l y i n t o tentacled arms which possess c i l i a t h a t d i r e c t food i n t o c i l i a t e d grooves which c a r r y i t t o the mouth. The alimentary canal i s U-shaped w i t h the anus outside the f r i n g e of tentacles. I n most species there i s only one p a i r o f pharyngeal s l i t s . Some pterobranchia are dioecious, but most are hermaphroditic. Although f e r t i l i z a t i o n i s external, the embryos g e n e r a l l y remain s h e l t e r e d w i t h i n the tubes. The sexually produced i n d i v i d u a l s then g i v e r i s e t o colonies by budding. These t i n y marine animals tend t o form l a r g e p l a n t Like colonies, whose i n d i v i d u a l s p r o j e c t Like small flowers a t the ends o f a branching s e r i e s o f tubes. The short body i s doubled-back on i t s e l f , so t h a t the anus opens a n t e r i o r l y over t h e back o f the head. I t i s generally believed t h a t they are a very p r i m i t i v e group rather than a degenerate hemichordate. Modern hemichordates appear t o be very s i m i l a r t o the f o s s i l g r a p t o l i t e s which were abundant i n the seas around the f i r s t f o s s i l evidence o f the vertebrates. THE LOWER CHORDATES (UROCHORDATES & CEPHALOCHORDATES) Small-sized marine animals e i t h e r being s e s s i l e ( t u n i c a t e s ) o r f r e e - l i v i n g (Lancelets). ORIGINS D i p l o b l a s t i c animals (u coelenterates) may be the most ancestral c o n d i t i o n since i t Lacks a middle Layer, the mesoderm. Higher animals a1 1 have a mesoderm and t r i p l o b l a s t i c . From the mesoderm muscular, s k e l e t a l and c i r c u l a t o r y systems are formed. There a r e two types of embryonic formation which allows f o r t h e development of t h i s Layer. I n echinoderms, chaetognaths, hemichordates and chordates the mesoderm a r i s e s from pouches growing outwards from the w a l l s o f the gut. These pouches remain i n the a d u l t as closed body c a v i t i e s . I n a l l other animals t h e mesoderm formation a r i s e s as s o l i d masses o f c e l l s budded o f f from an area near the p o s t e r i o r end o f the body, and the body c a v i t y a r i s e s by cleavage w i t h i n the mass o f the mesodermal c e l l s . This mesodermal development occurs i n molluscs and annelids, with the arthropods having a h i g h l y modified p a t t e r n o f development. A c l a d i s t i c c l a s s i f i c a t i o n would separate the echinoderms and chordates ( i n c l u d i n g hemichordates) as being the major d e v i a t i o n with the r e s t of the i n v e r t e b r a t e s developing independently. From t h i s theory the d i v i s i o n o f echinoderms and hemichordates from the other invertebrates i s r e f l e c t e d by t h e i r p a t t e r n o f cleavage, method of g a s t r u l a t i o n , and Larval development. This theory suggests t h a t chordates evolved d i r e c t l y from the most p r i m i t i v e metazoans, but t h i s would imply t h a t the middle body Layer, a t r u e body cavity, mouth and anus a l l evolved separately i n the echinoderms and protochordates and the other invertebrates. Resemblances between the lower chordates and same invertebrates have been the basis f o r several other theories o f chordate o r i g i n s . Annelid theory. Annelids and chordates are both b i L a t e r a l l y symnetrical and segmented, w i t h segmental excretory organs, a well-developed coelom, and Longitudinal blood vessels. I n v e r t i n g an annelid would place i t s nerve chord dorsal t o the d i g e s t i v e t r a c t , and the path o f blood flow would resemble t h a t o f chordates; however, the mouth would then be dorsal, u n l i k e a chordate, and other dorsoventral r e l a t i o n s would be altered. Annelids, moreover, have no s t r u c t u r e s suggestive of the notochord o r g i 11 s l i t s . Arthropod theory. Based on the steady progression o f forms from the simple invertebrates t o hunans as the most advanced L i f e forms. According t o the theory the arthropod-chordate theory, an arthropod became i n v e r t e d so t h a t the dorsal heart and v e n t r a l nerve cord became the chordate's v e n t r a l heart and dorsal nerve. The same problems o f t h i s theory e x i s t as t h a t f o r the annelid theory. Arachnid theory. Arachnids are thought t o have descended from an annelid ancestry. However, t h i s theory has the same problem o f the v e n t r a l nerve cord r e q u i r i n g the reversal o f dorsal and v e n t r a l surfaces. The segmentation i s s t i l l wrong, t h e r e i s no traces o f the notochord and i n t e r n a l g i l l s . The arthropod's j o i n t e d Legs could not have evolved i n t o the f i s h ' s fins. Echinoderm theory. s i m i l a r i t i e s o f the echinoderms t o the chordates include the indeterminate and r a d i a l cleavage. The t o r n a r i a Larva o f hemichordates and the b i l a t e r a l b i p i n n a r i a Larva o f echinoderms a r e both minute and transparent, w i t h almost i d e n t i c a l external 1 4 = c i l i a t e d bands and a Like nunber o f coelomic c a v i t i e s ; both a l s o having a dorsal pore. Even p r o t e i n s o f the blood serun are more c l o s e l y matched between hemichordates and echinoderms. Even the phosphorous compounds t h a t speed up the release of energy f o r muscle a c t i v i t y are s i m i l a r . A hypothetical sequence could be: Hemichordate Larva Echinoderm ( a u r i c u l a r i a ) Larva tunicate Larva * amphioxus + ostracoderm + + The problem i s t h a t the anatomy o f the L i v i n g echinoderms i s v e r y d i f f e r e n t t o hemichordates and protochordates. Ancestral echinoderms were, however, n e i t h e r r a d i a l l y symnetrical nor arranged i n groups o f f i v e nor where they a l l f r e e - l i v i n g . I n f a c t the p r i m i t i v e echinoderm Cothurnocystis appears t o be c l o s e l y r e l a t e d t o the pterobranchia. Garstangls theory f o r the o r i g i n o f the vertebrates = Branchiostoma (anphioxus) i s a small f i s h - l i k e marine animal (70 mn i n Length) occurring i n t r o p i c a l and temperate waters, and normally found p a r t l y b u r i e d i n sand. These animals are a l l f r e e swimning, possess a pronounced notochord, have major blood vessels t h a t are s i m i l a r t o t h a t of a f i s h , have m u l t i p l e g i l l s l i t s and a pharynx t h a t i s h i g h l y s p e c i a l i z e d f o r f i l t e r i n g . The g i l l s are therefore more o f a feeding r a t h e r than a r e s p i r a t i o n device. The o r i g i n o f t h e vertebrates i s about as u n c e r t a i n as the o r i g i n o f the chordates. For many years amphioxus was viewed as a L i v i n g r e l a t i v e f o r the vertebrate ancestors. I n a d d i t i o n t o having a l l the c h a r a c t e r i s t i c s o f t h e chordates, the a d u l t amphioxus also has resemblance t o the vertebrates. It i s generally agreed now t h a t the vertebrates are more l i k e l y t o have evolved from the other protochordate group, the Urochordates which include the t u n i c a t e seas q u i r t s ( r e d b a i t ) . Although the a d u l t i s a s e s s i l e animal t h a t i n no way appears t o resemble any other vertebrate, i t s f r e e swimning Larvae does. Garstang suggested t h a t t h i s urochordate acquired the a b i l i t y t o reproduce w h i l s t s t i l l a f r e e - l i v i n g Larvae and became the ancestor t o the vertebrates. The process o f a Larval stage being able t o reproduce i s c a l l e d paedomorphosis and i s not u n c m n i n chordates. Although Garstangls theory i s the most widely supported, there i s no r e a l evidence and a l t e r n a t i v e theories f o r v e r t e b r a t e e v o l u t i o n include d i r e c t descendance from the e x t i n c t echinoderm group Calcichordates. Another a l t e r n a t i v e i s based on the embryonic developnent o f vertebrates which has remained somewhat of a mystery; namely t h a t the embryonic o r i g i n s o f p a r t s o f the head, e s p e c i a l l y sense organs, muscles, and s t r u c t u r e s involved i n gas exchange, d i f f e r from those i n the r e s t o f the body. I t i s therefore suggested t h a t v e r t e b r a t e evolved from an amphioxus l i k e protochordate i n which the head gradually elaborated d u r i n g a t r a n s i t i o n from f i l t e r feeding t o a c t i v e predation. SUBPHYLUM UROCHORDATA: CLASS ASCIDIACEA s t o l o n i f e r a , Red-bait, p r e f e r s wave-beaten shores, presumably because the water movement aids the c i r c u l a t i o n o f water through i t s pharynx and ensures an adequate supply o f suspended food p a r t i c l e s . Not only does r e d - b a i t t h r i v e i n t u r b u l e n t waters, but the s i z e i t reaches i s d i r e c t l y r e l a t e d t o the amount of surge i t receives; i n c e r t a i n areas where wave a c t i o n i s p a r t i c u l a r l y strong, i t can reach 20 cm i n size. i s a s o l i t a r y tunicate, although they Live i n crowded groups each animal i s an independent u n i t . Each i n d i v i d u a l animal i s enclosed i n a t h i c k p r o t e c t i v e t e s t which i s attached t o the substrate by a holdfast. The t e s t has two dorsal t e a t - l i k e s t r u c t u r e s a t the apex which are the openings t o the buccal and a t r i a l siphons. Water i s drawn i n through the buccal c a v i t y and f i l t e r e d by the pharynx, and passes out through the a t r i a l siphon. The l a r v a i s f r e e swimning and s i m i l a r t o amphioxus. U n l i k e f i s h , emphioxus has no Lateral p a i r e d f i n s , b u t does have a dorsal f i n t h a t extends the e n t i r e Length of the body, a caudal f i n and a v e n t r a l f i n . A L L f i n s are strengthened by a dense connective t i s s u e t h a t forms f i n rays. The body w a l l s are d i v i d e d i n t o IoVooshaped muscle segments c a l l e d myotomes, which are separated from each other by connective tissue. They are arranged a l t e r n a t i v e l y on e i t h e r s i d e o f the body. Contraction of the myotomes produces Lateral body movements necessary f o r swimning. The protruding mouth i s surrounded by an o r a l hood from which p r o j e c t c i l i a t e d o r a l t e n t a c l e s ( c i r r i ) . The anus opens on the L e f t s i d e o f the body near t o the base o f the caudal f i n . The s k e l e t a l system i s composed o f a w e l l developed notochord, a rod of dense connective t i s s u e l y i n g dorsal t o the d i g e s t i v e t r a c t . Amphioxusls food consists o f minute organisms t h a t are f i l t e r fed i n t o the mouth by water c u r r e n t s produced by the c i l i a on the g i l l s . The d i g e s t i v e system consists o f a mouth which Leads t o a pharynx which has w a l l s pierced by 180 p a i r s o f g i l l s l i t s which sieve food from the water. A v e n t r a l c i l i a t e d groove, the endostyle secretes mucous. C i l i a conveys mucous up and over the g i l l bars and entraps food p a r t i c l e s . The c i l i a then pass the food-laden mucous i n t o the stomach. B l i n d hepatic cecae emerge from the stomach. There i s a s t r a i g h t i n t e s t i n e which Leads t o t h e anus. The c i r c u l a t o r y system i s s i m i l a r t o t h a t o f a f i s h , except t h a t i t Lacks the two charrrbered heart, but contains open sinuses. A s u b i n t e s t i n a l v e i n c o l l e c t s n u t r i e n t Laden blood from the i n t e s t i n e and c a r r i e s i t forward t o the hepaticp o r t a l v e i n and then t o the hepatic caecwn. The hepatic caecum Leads t o the v e n t r a l a o r t a which i s purped by rhythmical contractions i n t o t h e a f f e r e n t branchial a r t e r i e s t h a t supply the g i l l bars. I t then passes onto the e f f e r e n t branchial a r t e r i e s and onto the paired dorsal aortae. The blood i s oxygenated during the passage through the g i l l s and i s then taken back i n t o the dorsal aorta. The dorsal aorta d i v i d e s by way o f i n t e s t i n a l capi 1l a r i e s i n t o the s u b i n t e s t i n a l vein. The d i r e c t i o n o f the blood i s t h e r e f o r e p o s t e r i o r i n the dorsal vessels and a n t e r i o r i n t h e v e n t r a l vessels. Respiration occurs i n the pharynx which i s attached d o r s a l l y and hangs i n t o a c a v i t y c a l l e d the a t r i u m which i s Lined w i t h ectodermal epithelium. Water i s drawn through the mouth i n t o the pharynx and goes through the g i l l s l i t s i n t o the a t r i u n e x i t i n g through the atriopore. The g i l l s l i t s are framed by c i l i a t e d g i l l bars supported by rods o f dense connective tissue. The exchange of r e s p i r a t o r y gases occurs through the g i l l slits. Other UROCHORDATE CLASSES a r e LARVACEA and THALlACEA The excretory system i s composed o f s i n p l e nephridia that are s i t u a t e d d o r s a l l y - l a t e r a l l y . Each bears c l u s t e r s o f f l a g e l l a t e d flame c e l l s . There are about 100 nephridia connected t o the d o r s a l coelom w i t h i n the a t r i a l cavity. CEPHALOCHORDATA EXAMPLE BRANCHIOSTOMA (AMPHIOXUS) The nervous system consists o f a c e n t r a l nerve cord t h a t runs dorsal t o t h e d i g e s t i v e t r a c t and r e s t i n g on the notochord. The c e n t r a l canal transverses t h e e n t i r e Length and widens a t t h e a n t e r i o r end t o form a cerebral v e s i c l e ( l o c a t i o n o f t h e brain). Two p a i r s o f sensory nerves a r i s e from the v e s i c l e and serve t h e a n t e r i o r end. The r e s t o f t h e nerve cord gives o f f t h e nerves on the opposite sides t h a t a l t e r n a t e with each other. These nerves are o f two kinds, the dorsal nerves with a sensory f u n c t i o n and t h e v e n t r a l nerves w i t h a motor function. Sense organs include: 1) O l f a c t o r y p i t 2) Epidermal sensory & l l s on the o r a l and v e l a r tentacles 3) Rows o f simple eyes ( 1 ganglion and 1 pigment c e l l ) The reproductive system i s composed o f p a i r e d gonads t h a t p r o j e c t i n t o t h e atrium. The sexes are separate and the egg and sperm c e l l s are discharged i n t o the a t r i a l c a v i t y and reach t h e e x t e r i o r v i a an atriopore. F e r t i l i z a t i o n i s external. The r e s t o f t h e chordates, c o l l e c t i v e l y r e f e r r e d t o as the vertebrata a l l have an enlarged b r a i n enclosed i n a b r a i n case, o r craniun, and a segmented spinal c o l m o f vertebrae t h a t become t h e a x i a l support o f t h e body. T y p i c a l l y t h e body comprises a head, neck, trunk, and a t a i l . These classes show a progressive series o f s t r u c t u r a l and f u n c t i o n a l advances i n a l l organ systems besides the features o f the notochord and g i l l s l i t s . 1) The body covering i s a s t r a t i f i e d epithelium and dermis, w i t h many mucous glands i n aquatic species; most f i s h e s are covered w i t h p r o t e c t i v e scales; the e x t e r i o r i s c o r n i f i e d on Land dwellers, with scales on r e p t i l e s , feathers on b i r d s and h a i r on mamnals; feathers and h a i r form i n s u l a t e d body coverings. 2) The i n t e r n a l and j o i n t e d skeleton i s o f c a r t i l a g e i n Lower vertebrates and o f bone i n higher groups; i t supports and p r o t e c t s various organs; the cranium s h e l t e r s t h e b r a i n and has p a i r e d capsules t o contain organs o f special sense; a s e r i e s o f v i s c e r a l arches supports t h e g i l l region, and c e r t a i n arches become the jaws and other s t r u c t u r e s o f t h e head region; t h e v e r t e b r a l column extends from the base o f the cranium t o the end o f t h e t a i l and has neural arches d o r s a l l y t o house t h e nerve cord. Two p a i r s o f appendages, the f i n s o f f i s h e s and Limbs o f tetrapods, w i t h j o i n t e d s k e l e t a l supports, are a r t i c u l a t e d with the v e r t e b r a l c o l m through Limb g i r d l e s . 3) On t h e skeleton are muscles t h a t move i t s p a r t s and provide f o r Locomotion. 4) The Long d i g e s t i v e t r a c t i s v e n t r a l t o the v e r t e b r a l c o l m ; the mouth contains a tongue and u s u a l l y teeth; the anus opens a t the end o f the trunk; the Liver and pancreas are two Large d i g e s t i v e glands that pour t h e i r secretions through ducts joined t o t h e intestine. 5 ) The c i r c u l a t o r y system includes a well-developed muscular heart o f two, three, o r four chambers, Located v e n t r a l l y t o t h e d i g e s t i v e t r a c t ; i t s contractions propel the blood through a closed system of arteries, c a p i l l a r i e s , and veins, the flow being a n t e r i o r l y on the v e n t r a l s i d e and p o s t e r i o r l y i n the dorsal arteries; t h e blood plasma contains both white and red corpuscles, t h e L a t t e r w i t h haemoglobin as a r e s p i r a t o r y pigment; a system o f Lymph vessels i s present; p a i r e d a o r t i c arches transport blood from t h e heart t o t h e g i l l s i n Lower vertebrates; progressive separation o f t h e r e s p i r a t i o n (pulmonary) and systemic blood paths through t h e heart Leads t o r e a l i z a t i o n of regulated body temperature i n t h e warm blooded (homeothermat) b i r d s and mamnals. 6) Respiration i n the Lower forms i s by p a i r e d g i l l s ; t e r r e s t r i a l species have Lungs developed from outpocketings o f the d i g e s t i v e t r a c t . 7) The paired excretory organs, o r kidneys, discharge through duct openings near o r through t h e anus; i n lower forms, the organs are o f segmental nature and d r a i n wastes from both the coelom and t h e blood; i n higher forms they are non-segmental and d r a i n o n l y from the blood; a bladder f o r storage o f u r i n e occurs i n many. 8) The b r a i n becomes r e g i o n a l l y d i f f e r e n t i a t e d as t o s t r u c t u r e and function; the cerebral hemispheres and cerebellum enlarge, e s p e c i a l l y i n higher forms; there are 10 o r 12 p a i r s o f c r a n i a l nerves i n t h e head t h a t serve both motor and sensory function, i n c l u d i n g t h e paired organs of special senses (smell, sight, and hearing plus e q u i l i b r a t i o n ) ; from the nerve cord a p a i r of spinal nerves serves each p r i m i t i v e body somite; an autonomic nervous system regulates i n v o l u n t a r y functions o f i n t e r n a l organs. 9) A series o f endocrine glands (thyroid, p i t u i t a r y , e t c ) provides hormones, transported by t h e blood stream, t h a t regulate b o d i l y processes, growth, and reproduction. 10) With r a r e exceptions t h e sexes are separate, and each has a p a i r o f gonads t h a t discharge sex c e l l s through ducts opening i n t o o r near t h e anus. SUBPHYLUM VERTEBRATA: CLASS AGNATHA Two Living representative, the Lamprey (order Petromyzontida and Hagfish (phylogenetic r e l a t i o n s h i p s are unclear but has been put i n t o the order Myxinida). Lampreys are both marine and fresh water, whereas hagfish are e x c l u s i v e l y marine. These Lowest o f Living vertebrates Lack t r u e jaws, and t h e i r nearest a l l i e s are the ancient ostracoderms o f t h e S i l u r i a n and Devonian times. The Lamprey-hagfish Lines o f e v o l u t i o n diverged e a r l y and probably had separate o r i g i n s among the ostracoderm ancestors. Lampreys and hagfish are p r i m i t i v e animals because they Lack paired f i n s , they posses p o o r l y developed c a r t i l a g i n o u s skeletons, they have a s i n g l e n o s t r i l and they Lack jaws. Both Lampreys and hagfish are h i g h l y modified ( p a r t i c u l a r l y i n the v i c i n i t y o f t h e mouthparts) f o r p a r a s i t i z i n g (Lamprey) and scavenging (hagfish). The ancestral form however, was probably f i l t e r - f e e d i n g Ostracoderms. OSTRACODERMS These animals were f i r s t found i n t h e mid-Ordovician (500 t o 450 m i l l i o n years ago). Their c h a r a c t e r i s t i c s include: A s i n g l e dorsal n o s t r i l They were without p a i r e d appendages ( f i n s ) , but some had peculiar f l a p s p r o j e c t i n g from behind t h e head. ALL had a bony armour o r scales and i n some t h e head contained an i n t e r n a l skeleton of bone. This suggests t h a t bone i s an archaic feature d a t i n g t o t h e e v o l u t i o n o f the f i r s t vertebrates. The bony armour may have provided p r o t e c t i o n against ancient water scorpions (e.g.Pterygotus). Other suggestions are t h a t t h e armour prevented undue water Loss o r represented a store of calciun salts. L i k e a l l agnathans, they Lacked jaws, however, t h e i r pharynx possessed paired g i l l s l i t s . They possessed a dorsal s p i n a l cord, and notochord. ; ' The head region o f some possessed a pronounced dorsal Land Lateral sensory f i e l d s Leading t o speculation t h a t @' they could generate e l e c t r i c f i e l d s . - = They evolved i n t o a great nunber o f forms varying from f l a t bottom-dwelling animals t o a c t i v e f a s t swimning animals (Birkenia). Some forms even Lost t h e i r bony p l a t e s and became covered w i t h minute d e n t i c l e s (Thelodus). opening . NERVOUS SYSTEM The b r a i n i s p r i m i t i v e , dominated by f o r e b r a i n w i t h a Large p a i r o f o l f a c t o r y Lobes. The cerebral hemispheres are attached t o a diencephalon, and there i s a v e n t r a l l y attached broad infund.ibulun and above i t a pineal structure. The midbrain possesses a l a r g e p a i r o f o p t i c Lobes, and t h e h i n d b r a i n has rudimentary cerebellun. There are ten p a i r s o f c r a n i a l nerves. LAMPREY (ORDER PETROMYZONTIDA) SENSE ORGANS EXTERNAL FEATURES * Body i s c y l i n d r i c a l w i t h a L a t e r a l l y compressed t a i l possessing a caudal f i n . The s k i n i s without scales, but has an epidermal gland which produces the slimy secretions. There i s a row o f sensory p i t s forming a Lateral Line. The mouth i s h i g h l y modified and consists o f s u c t o r i a l disc, a mouth t h a t i s h e l d open by a r i n g o f c a r t i l a g e , bordered by sensory p a p i l l a e and horny l1teethl1 Like structures. From t h e mouth protrudes a p i s t o n - l i k e tongue. There are a p a i r o f eyes, s i n g l e nasal opening and seven g i l l s l i t s . The anus i s on t h e v e n t r a l surface j u s t behind the urogenital opening. SKELETAL SYSTEM A notochord p e r s i s t s throughout. The a x i a l skeleton i s composed o f small c a r t i l a g i n o u s neural arches, with the head supported and protected by a c a r t i l a g e s k u l l . A c a r t i l a g e branchial basket p r o t e c t s t h e g i l l s . There i s stout c a r t i l a g e i n the tongue. MUSCULAR SYSTEM Short segmented muscles which appear t o have a IgZgg shape i n s i d e view. A l t e r n a t e contractions produce rhythmic Lateral movement o f t h e f l a n k s which propel the animal forward. Radial muscles surround the mouth and r e t r a c t o r muscles occur i n t h e tongue. DIGESTIVE SYSTEM Oral d i s c and i t s horny t e e t h attach the Lamprey t o other f i s h . The tongue rasps i n t o the f l e s h o f the f i s h using i t s r e t r a c t o r muscles. The d i g e s t i v e system consists o f an oesophagus, a v e n t r a l pharynx, no d i f f e r e n t i a t e d stomach, a Long s t r a i t i n t e s t i n e which has i n t e r n a l s p i r a l l y arranged f o l d s (typhlosole) which increases surface area f o r absorption. The d i g e s t i v e system terminates i n an anus. A Liver i s present i n association w i t h t h e d i g e s t i v e system. CIRCULATORY SYSTEM Consists o f heart, a r t e r i e s , veins and Lymphatic sinuses. The heart Lies i n the p e r i c a r d i a l c a v i t y and consists o f an a t r i u n which receives blood and v e n t r i c l e which pumps blood. RESPIRATORY SYSTEM Seven p a i r s o f g i l l pouches which open i n t e r n a l l y i n t o the pharynx and e x t e r n a l l y v i a the g i l l s l i t s . There are numerous g i l l filaments supplied w i t h c a p i l l a r i e s . Gaseous exchange occurs as water passes through the g i l l pouches. Water i s drawn i n t o the g i l l pouch through t h e g i l l s l i t s (an adaptation t o a p a r a s i t i c L i f e r e q u i r i n g t h a t the mouth be attached t o a host). The Larval Lamprey ( c a l l e d an amnocoete) i s not p a r a s i t i c , and water passes through t h e mouth and over the g i l l s ( t h e p a t t e r n t h a t occurs i n bony f i s h ) . EXCRETORY SYSTEM Linked with t h e reproductive system (= urogenital system). The kidneys L i e along t h e dorsal wall o f the body. U r i n e passes v i a a u r i n a r y duct i n t o the g e n i t a l sinus which opens t o the e x t e r i o r v i a a urogenital The o l f a c t o r y sac i s contained i n the nasal capsule and i s connected t o the outside through t h e s i n g l e nasal opening occurring on the dorsal surface. There i s a balancing organ which l i e s i n the a u d i t o r y capsule and consists o f two semicircular canals. The eyes are p r i m i t i v e , yet allow good vision. P o s t e r i o r t o the nasal opening i s t h e pineal eye w i t h a Lens and a pigmented r e t i n a . The pigments o f t h e eye vary with the Lamprey switching from Rhodopsin under marine conditions t o Porphyropsin under freshwater conditions. Lampreys are also able t o produce e l e c t r i c a l f i e l d s . REPRODUCTION The sexes are separate, with t h e gonad being protandric, t h a t i s f i r s t producing male gametes and then female gametes w i t h i n t h e same i n d i v i d u a l . A s i n g l e Large Long gonad f i l l s t h e abdominal cavity. There are no g e n i t a l ducts, w i t h egg o r sperm being discharged i n t o abdominal c a v i t y and passes through paired g e n i t a l pores i n t o t h e u r o g e n i t a l sinus and t h e passes outside the body external f e r t i l i z a t i o n takes place. Spawning always occurs i n freshwater, w i t h a female lamprey producing up t o 100 000 eggs. A f t e r spawning the female Lampreys die. The eggs hatch i n t o a Larvae c a l l e d t h e amnocoete and remain i n t h i s condition f o r 3 t o 12 years. The Larva obtains i t s food through i n s p i r a t i o n o f water and using mucous t o entangle t h e food and pass i t t o t h e small i n t e s t i n e ( i n a very s i m i l a r fashion t o t h a t o f amphioxus). The amnocoete Larva undergoes metamorphosis t o become an a d u l t Lamprey and t h e returns t o t h e sea. The lamprey d i f f e r s from t h e hagfish which has a mouth t h a t i s surrounded by sensory tentacles and a l s o has a varying nunber o f g i l l pouches ( f i v e t o f i f t e e n ) . CLASS PLACOOERMI These are Large, h e a v i l y armoured f i s h possessing jaws, and were possibly the f i r s t vertebrates t o possess them ( t h i s may, however, be i n dispute). The r i s e o f the placoderms probably c o n t r i b u t e d t o t h e e x t i n c t i o n o f the ostracoderms. These f i s h a l s o characterized the f i r s t attempts a t developing p a i r e d limbs. There were many experiments with appendages, although many placoderms were s t i l l without f i n s , but s t i l l possessed Large hollow f i x e d spines p r o j e c t i n g outwards from the shoulder regions. Most placoderms were predatory f i s h and some reached 10 metres i n length. The most c h a r a c t e r i s t i c feature are the heavy armour p l a t e s on the outside, e s p e c i a l l y around t h e head. I n some o f these bony p l a t e s served t h e f u n c t i o n o f jaws and teeth, o r even Large crushing plates. The placoderms were f a r removed from the main evolutionary Line o f the vertebrates. Therefore, few, i f any, o f t h e known types can be considered ancestral t o Later vertebrates. An ancestral form may have Lost i t s p l a t e s o f bony armour, and could have given r i s e t o the sharks and chimaeras. - The appendicular skeleton includes a V shaped pectoral g i r d l e supporting t h e pectoral f i n s , a f l a t t e r p e l v i c g i r d l e w i t h f i n s and a smaller j o i n t e d c a r t i l a g e which provides i n t e r n a l support f o r the Lateral f i n s . CLASS CHONDRICHTHYES (Cartilaginous f i s h ) CURRENTLY THE MOST PRIMITIVE VERTEBRATES UHICH POSSESS t Complete and separate vertebrates t Movable jaws MUSCULAR SYSTEM t Paired appendages Body and t a i l muscles segmented ADVANCES OVER THE CLASS AGNATHA t Scales (placoid) covering the body t Two p a i r s o f Lateral f i n s t t Movable jaws which a r t i c u l a t e w i t h the craniun Enamel covered teeth t Three semicircular canals DIGESTIVE SYSTEM MWTH Teeth derived from modified p l a c o i d scales, replacement teeth develop continuously F l a t tongue adhere t o the f l o o r o f the mouth. On the sides o f t h e wide pharynx are openings Leading t o separate g i l l s l i t s and spiracles. Short oesophagus leading t o a "Jl1 shaped stomach which ends w i t h a c i r c u l a r sphincter muscle ( p y l o r i c valve) I n t e s t i n e f o l l o w s and i s connected t o a cloaca and anus. I n t e s t i n e contains a s p i r a l valve, which i s covered by a mucous membrane which slows down the food and increases absorptive surfaces. L i v e r has two Long Lobes, secretes b i l e v i a a duct and a conspicuous g a l l bladder exists. Pancreas Lies between t h e stomach and the intestine, i t s duct j o i n i n g the l a t t e r j u s t below the b i l e duct. A r e c t a l gland attaches d o r s a l l y a t the junction o f the i n t e s t i n e and the cloaca and removes excess s a l t from t h e blood. Paired reproductive organs LESS ADVANCED THAN A BONY FISH (CLASS OSTEICHTHYES) SINCE t Skeleton i s made o n l y o f c a r t i l a g e t P l a c o i d scales o n l y t Separate g i l l c l e f t s t Possesses a p a i r o f s p i r a c l e s connected t o the pharynx t Has no swim bladder COELOM (Body Cavity) EXAMPLE IS THE DOGFISH (SQUALUS) t Stomach, intestine, and other i n t e r n a l organs l i e i n the Large body c a v i t y o r coelom. t I t i s l i n e d w i t h a smooth membrane: t h e peritoneun and t h i s covers the organs. t The peritoneum i s supported from the mid dorsal wall o f the coelom by t h i n mesenteries. t The transverse septun separates coelom from the c a v i t y containing t h e heart. EXTERNAL FEATURES t t t t t t t Blunt pointed head Trunk s p i n d l e shaped Two separate median dorsal f i n s A median caudal f i n Two p a i r s o f l a t e r a l f i n s (pectoral and p e l v i c ) Mature males have a clasper on t h e i r p e l v i c fins Anal f i n CIRCULATORY SYSTEM HEAD t t t t t Two n o s t r i l s L a t e r a l eyes without e y e l i d s Five oval g i l l s l i t s open a n t e r i o r l y t o each pectoral f i n S p i r a c l e ( g i l l c l e f t ) opens behind each eye The heart i s beneath t h e g i l l region: consists o f a t h i n - w a l l e d dorsal sinus venosa t h a t receives blood, followed by t h e a t r i u n , then the t h i c k walled v e n t r i c l e and t h e conus arteriosus. Blood passes a n t e r i o r l y i n t o t h e v e n t r a l aorta where f i v e p a i r s o f a f f e r e n t branchial a r t e r i e s d i s t r i b u t e t o the c a p i l l a r i e s i n t h e g i l l s f o r aeration, four p a i r s o f e f f e r e n t branchial a r t e r i e s c o l l e c t blood i n t o the dorsal aorta which extends along mid-dorsal w a l l o f the coelom. BODY COVERING Diagonal rows o f minute p l a c o i d scales, each backward p o i n t i n g and covered wi.th enamel over a base p l a t e o f dentine. SKELETON P r i n c i p a l a r t e r i e s are: t C a r t i l a g e r e i n f o r c e d w i t h limy deposits t A x i a l skeletons: s k u l l / v e r t e b r a l c o l m , w i t h each vertebra having a spool shaped centrun, concaved a t both ends, a neural arch housing t h e nerve cord. The notochord p e r s i s t s i n spaces between t h e vertebrae. t t t b t t The s k u l l i s composed o f a cranium housing the brain, possesses p a i r e d capsules f o r olfactory, o p t i c and a u d i t o r y organs. The v i s c e r a l skeleton consists o f jaws, the hyoid arch and f i v e p a i r s o f branchial arches supporting the g i l l region. t Paired e x t e r n a l / i n t e r n a l c a r o t i d s -. head Paired subclavians -. p e c t o r a l f i n s Coelic -. stomach, l i v e r , i n t e s t i n e Anterior mesenteric -. spleen and p o s t e r i o r intestine Posterior mesenteric -, t o r e c t a l gland Renal and gonadic -. t o kidneys/reproductive organs Paired i l i a c s -. p e l v i c f i n s MIDBRAIN P r i n c i p a l veins F b b b b Paired renal p o r t a l kidneys Paired postcardinal + p a r a l l e l i n g the kidneys Paired Lateral abdominal veins e i t h e r side o f t h e body c a v i t y Jugular and a n t e r i o r cardinal veins (sinuses) r e t u r n blood from the head region Blood from t h e d i g e s t i v e t r a c t flows v i a the hepatic p o r t a l v e i n and i s f i l t e r e d through the l i v e r and then i s c o l l e c t e d i n the hepatic veins which are connected t o the sinus venosus. + + RESPIRATORY SYSTEM F By opening and c l o s i n g t h e mouth, sharks draw i n water and passes i t out through t h e g i l l s l i t s and spiracles. b Two rounded Lobes d o r s a l l y s i t u a t e d HINDBRAIN b b F b b Large median dorsal cerebellun over t h e opentopped medulla oblongata Eleven c r a n i a l nerves (one t o t e n p l u s zero) Spinal nerve protected by vertebrae Paired spinal nerves t o each body segment Sympathetic nervous system: s e r i e s o f ganglia, roughly segmental above t h e postcardinal veins. REPRODUCTIVE SYSTEM SEXES ARE SEPARATE The g i l l s are composed o f many p a r a l l e l , slender filaments t h a t contain c a p i l l a r i e s . Male sperm develops i n long t e s t e s From each testes vasa deferens runs p o s t e r i o r l y on the v e n t r a l surface o f t h e kidney and empties i n t o the u r o g e n i t a l p a p i l l a . Blood from the v e n t r a l aorta passes through these c a p i l l a r i e s , discharges carbon dioxide and absorbs oxygen dissolved i n water, then continues i n t h e dorsal aorta. At mating claspers are placed close together, and they are inserted i n t o t h e cloaca o f the females. Seminal f l u i d flows down the canal formed by t h e adjacent clasper grooves. EXCRETORY SYSTEM b The two slender kidneys L i e imnediately above the coelom on e i t h e r s i d e o f t h e dorsal aorta. b U r i n e from each kidney i s c o l l e c t e d i n a series o f segmental tubules t h a t j o i n a l o n g i t u d i n a l duct, t h e ureter, Leading p o s t e r i o r l y ; t h e two u r e t e r s empty through a s i n g l e urogenital p a p i l l a , Located d o r s a l l y i n the cloaca. b Females have two l a r g e ovaries, two oviducts extend the length of t h e body cavity, t h e i r a n t e r i o r ends j o i n i n a s i n g l e l a r g e funnel through which the eggs enter. The forward p a r t o f the each duct i s d i l a t e d as a s h e l l gland, and i n ovoviviparous species, Like the dog f i s h the p o s t e r i o r p a r t i s enlarged as a 88uterus88 to contain the young during development. The oviducts open separately i n t o the cloaca. CLASSIFICATION OF CHONDRICHTHYES . SENSE ORGANS b Two n o s t r i l s ( o l f a c t o r y sacs) b Pharynx: scattered t a s t e buds b Eyes: b b I1Ear" Lateral Line moved by three p a i r s o f muscles attaching t h e eyeball t o i t s socket and are without l i d s . Retina composed o f rods only, therefore no colour v i s i o n i s possible. Organ o f ba 1ance Three s e m i - c i r c u l a r canals A f i n e groove along each s i d e o f t h e trunk and Line t a i l . Contains a slender canal w i t h many small openings t o the surface. W i t h i n t h e canal are sensory h a i r s connecting t o a branch o f t h e tenth c r a n i a l nerve. They respond t o Low frequency pressure s t i m u l i i n t h e surrounding water. On the head are other sensory canals opening by senses pores; each pore Leads t o a small chamber (Ampulla o f Lorenzini) containing an electroreceptor w i t h sensory h a i r s connecting t o nerve fibres. SUBCLASS ELASMOBRANCHII ORDER CLADOSELACHIDA ORDER XENACANTHIDA E a r l y f r e s h water sharks * Extinct * ORDER GALEOMORPHA L i v i n g sharks Mollusc feeding forms Nurse sharks Modern sharks (great whites) Tiger sharks S i x - g i l l e d sharks ORDER SQUALOMORPHA Dogfish and small r e l a t e d sharks ORDER BATOIDEA Skates and rays; Ibbottom L i v i n g formsbb flattened dorsoventral ly, pectoral f i n s enlarged and Locomotion i s achieved through undulations, Lost the caudal fins; no Lateral thrashing o f the t a i l . Mouth on bottom. Water brought i n v i a s p i r a c l e s on dorsal surface (behind the eyes) NERVWS SYSTEM More advanced than the lamprey. FOREBRAIN b F b b b Two o l f a c t o r y sacs Large o l f a c t o r y t r a c t s + o l f a c t o r y Lobe Paired cerebral hemispheres on t h e diencephalon Dorsal p i n e a l body Ventral infundibulun and hypophysis Ancestral sharks, appeared i n t h e Devonian. Paired f i n s but no claspers. * Extinct * SUBCLASS HOLOCEPALI S h a r k - l i k e but w i t h crushing tooth plates. Most forms are extinct. ORDER CHIMAERIDA Chimaeras o r r a t f i s h , Flap o f s k i n covering the separate branchial slits. w SPINY SHARKS CLASS ACANTHODIANS Heart i s two chambered (one a t r i u n and one v e n t r i c l e ) . I n association w i t h t h e heart i s a sinus venosa and conus arteriosus. Heart contains only venous blood; red blood c e l l s are nucleated and oval. These may w e l l have been t h e e a r l i e s t gnathostomes (vertebrates w i t h t r u e jaws). They were small f i s h which s u p e r f i c i a l l y resembled sharks. While the contemporary placoderms a l s o had jaws, the acanthodian jaw i s b a s i c a l l y t h e same as t h a t o f the bony f i s h (Class Osteichthyes). I t i s also thought t h a t they had Lungs with which they used f o r e i t h e r breathing or as h y d r o s t a t i c organs ( f u n c t i o n a l swim bladders?). - The e a r l y acanthodians are t y p i f i e d by the genus Climatius which was f i r s t found i n the Late S i l u r i a n period. They were small, o n l y a few centimetres i n Length. Their bodies are t y p i c a l l y f i s h - l i k e (spindleshaped) w i t h t h e p o s t e r i o r body being t i l t e d upwards, with the caudal f i n being underneath c r e a t i n g a t y p i c a l heterocercal t a i l . There were two Large t r i a n g u l a r dorsal median f i n s , f i v e p a i r e d v e n t r a l median f i n s as well as p a i r e d pectoral and p e l v i c f i n s . Climatius was protected by a dermal armour o f small rhombic o r diamond-shaped scales which covered the e n t i r e body and continued over t h e head i n t h e form o f r e g u l a r l y arranged plates. These f i s h were characterized by extremely Large eyes r e l a t i v e t o t h e i r body s i z e and eyes t h a t are placed f a r forward. I n contrast the nasal region f o r o l f a c t o r y senses were rather poorly developed. The upper jaw was generally o s s i f i e d and well developed. On the sides o f the heads were the g i l l coverings o r opercular flaps, one t o each o f the f i v e g i 11 arches, a d over these there were larger opercular coverings o f s t i f f bony rods. Although Climatius represents a p r i m i t i v e ancestral form the acanthodians d i v e r s i f i e d i n the same ways t h a t the bony f i s h have; namely Long slender e e l - l i k e forms t o deep-bodied f a s t swimning forms. I n some there was a Loss o f t h e p a i r e d f i n s and a tremendous exaggeration o f the dorsal f i n spines. Skeleton c h i e f l y o f bone ( c a r t i l a g e i n Vertebrae a r e many and p r i m i t i v e sturgeons). d i s t i n c t ; t a i l u s u a l l y homocercal (even-sized above and below caudal vertebrae). Primitive f i s h o f t e n have heterocercal t a i l s (caudal vertebrae bend upwards) R e l i c t s o f a notochord many s t i l l p e r s i s t . Respiration i s by p a i r s o f g i l l s on bony g i l l arches contained w i t h i n a comnon chamber on each side o f the pharynx and covered by a bony operculun. Usually possess a swim bladder which sometimes has a duct connecting t o t h e pharynx. I n some f i s h t h e swim bladder i s a Lungfish), t r u e Lung (Dipnoi f i s h : i n d i c a t i n g t h a t t h e e a r l i e s t f i s h had Lungs. b b Cranial nerves zero t o t e n are present. Body temperature dependent on envirorment and behaviour ( p o i k i lothermal) may be elevated i n Large a c t i v e f i s h . Gonads t y p i c a l l y paired; u s u a l l y oviparous (some ovovivaparous and viviparous forms), f e r t i l i z a t i o n u s u a l l y external (exceptions exist). Eggs are u s u a l l y minute but can be Large such as t h e 100 mn eggs o f t h e Coelacanth. PHYLOGENY OF OSTEICHTHYES Divided i n t o SUBCLASS SARCOPTERYGII Divided i n t o Acanthodian were most abundant i n the Devonian and thereafter d e c l i n e d and were e x t i n c t by the end o f the Palaeozoic. The phylogenetic r e l a t i o n s h i p o f these f i s h t o the other jawed f i s h e s (placoderms, c a r t i l a g i n o u s and bony f i s h ) i s unclear. I t i s now generally excepted that they belong t o a class o f t h e i r own, but they have s i g n i f i c a n t r e l a t i o n s h i p s w i t h osteichthyes (* opercular flaps, jaws and p a i r e d f i n s ) . BONY FISH CLASS OSTEICHTHYES CHARACTERISTICS b Skin w i t h many mucous glands, u s u a l l y w i t h embedded bony dermal scales (Ganoid i n p r i m i t i v e forms, otherwise c y c l o i d o r ctenoid); some f i s h have Lost t h e i r scales; both median and p a i r e d f i n s are present (some exceptions and modifications), f i n s are supported by f i n rays composed o f c a r t i l a g e o r bone; no Limbs. Mouth u s u a l l y terminal and possessing teeth; jaws w e l l developed and a r t i c u l a t e d t o the s k u l l ; two o l f a c t o r y sacs t h a t are s i t u a t e d d o r s a l l y and are u s u a l l y not connected t o the mouth cavity; eyes are u s u a l l y w e l l developed possessing rods and cones ( t h e L a t t e r f o r colour v i s i o n ) and not possessing eyelids. F i n - r a y f i s h = SUBCLASS ACTINOPTERYGII Lobe-finned = SUBCLASS SARCOPTERYGII ORDER CROSSOPTERYGII SUBORDER COELACANTHINI SUBORDER RHIPIDISTIA (These were t h e ancestors t o t h e tetrapods) ORDER DIPNOI (Lungfish) CROSSOPTERYGII These were the most comnon f i s h i n t h e Devonian times. Generally these f i s h were aggressive, predacious f i s h , t h a t were ancestral t o the tetrapods (amphibian and above groups). They had already become r a r e by the Carboniferous times and w i t h one exception were e x t i n c t by the close o f the Palaeozoic. The coelacanths are stub-nosed. small iawed f i s h which although once though t o be e x t i n c t are-now known t o be the only Living Crossopterygian species. I n 1939 a Coelacanth species was found o f the coast o f South A f r i c a (East London). This species has been c l a s s i f i e d i n the genus Latimeria and represents the closest f i s h r e l a t i v e t o the tetrapods (four-Legged animals). Latimeria i s a h i g h l y specialized deep sea f i s h , whose d i s t a n t ancestors would have evolved towards t h e tetrapod structure. Coelacanths are very u n l i k e the Crossopterygians o f t h e Devonian times t h a t evolved towards the amphibian condition. These Devonian Crossopterygians possessed a s k u l l which had a passage that Linked the n o s t r i l w i t h the roof o f i t s mouth. ALL tetrapods have t h i s feature and confirm t h a t these f i s h were on the evolutionary Line t h a t Lead t o the f i r s t Land vertebrates ( t h e amphi bians represented by an animal c a l l e d Eusthenopteron). . good o l f a c t o r y senses and a L a t e r a l l i n e system. COELACANTH CHARACTERISTICS b b Heavy cosmoid scales b Fleshy f i n s b Lungs a r e Large s a c - l i k e and f i l l e d w i t h f a t and connective t i s s u e . b Bone has been reduced and replaced w i t h cartilage. b EVOLUTIONARY HISTORY OF THE ACTINOPTERYGIANS Lack i n t e r n a l n o s t r i l s , t h e nasal sacs a r e f i l l e d w i t h j e l l y , t h e f u n c t i o n o f which i s unknown. I s o l a t e d scales were found i n t h e l a t e S i l u r i a n , w h i l e the f i r s t body f o s s i l s were found i n t h e Middle Devonian. The e a r l y f i s h were small, possessing heavy scales and complex, f u l l y o s s i f i e d s k u l l s , characterized by several c i r c u n - o r b i t a l bones. Toothbearing bones characterize both t h e upper and Lower jaws. The eyes were extremely Large, and t h e mouths were Long. These f i s h were c h a r a c t e r i z e d by a heterocercal t a i 1 (possessing an extremely Long upper Lobe), a s i n g l e dorsal f i n was s i t u a t e d f a r back on the body. The v e n t r a l surface had an anal f i n a l s o s i t u a t e d f a r back on t h e body. P e c t o r a l and p e l v i c f i n s were widely separated. The vertebrae were incompletely o s s i f i e d and t h e notochord was v e r y conspicuous. Heart i s a simple s e r i e s o f Linear chambers. t B r a i n case i s Large, but t h e b r a i n w i t h i n i s v e r y small. w No p i n e a l foramen b Loss o f s a l t s i s prevented by maintaining a h i g h osmotic concentration achieved through r e t e n t i o n o f urea i n t h e blood (a s i m i l a r mechanism used by sharks). b Eggs a r e Large (100 mn) and develop w i t h i n t h e maternal oviduct (ovoviviparous) The actinopterygian f i s h were t r a d i t i o n a l l y d i v i d e d i n t o three groups. Although these groups appear t o Lack s c i e n t i f i c i n t e g r i t y ( a c l a d i s t would c e r t a i n l y not recognize them) they do represent stages i n t h e e v o l u t i o n o f t h i s f i s h group, and t h e r e f o r e I have r e t a i n t h i s somewhat o l d fashion scheme. The groups are: b CHONDROSTEI (most p r i m i t i v e forms) b HOLOSTE I ( i n t e r m e d i a t e forms) ( = NEOPTERYGII) b TELEOSTEI ORDER DIPNOI (Lungfish) - Three L i v i n g genera. One genera o c c u r r i n g i n South America, South A f r i c a and A u s t r a l i a respectively. They Like t h e Coelacanths share s i m i l a r anatomical features w i t h t h e amphibians. CHONDROSTEI These were abundant i n t h e Palaeozoic and a r e sometimes r e f e r r e d t o as Palaeoniscoids. They were g e n e r a l l y small f i s h w i t h u p t i l t e d (heterocercal) t a i l s and scales covered w i t h ganine ( = ganoid scales). DIPNOI CHARACTERISTICS b Cosmoid scales b Fleshy f i n s b Reduction i n o s s i f i c a t i o n o f bone b Functional Lungs b A u s t r a l i a n forms can s u r v i v e i n stagnant water by a i r - b r e a t h i n g . The other genera can s u r v i v e complete d r y i n g up o f the water by burrowing and a e s t i v a t i n g (dormancy phase). The A f r i c a n Lungfish a r e so dependent on a i r t h a t i f kept underwater they w i l l drown. (most advanced forms) A great v a r i e t y o f forms e x i s t e d i n t h e Palaeozoic, ranging from Long t h i n almost e e l - l i k e forms t o L a t e r a l l y compressed deep-bodied forms (resemble angel f i s h ) . The group i s g e n e r a l l y e x t i n c t except f o r the f o l l o w i n g examples. b SUBCLASS ACTINOPTERYGII Sturgeon f i s h jAcipenser) represented by s i x t e e n species. These have g e n e r a l l y Lost t h e i r ganoid scales except i n t h e t a i l r e g i o n and t h e r e have a naked skin. They have replaced scales w i t h a p a r t i a l covering o f p l a i n bony plates. The skeleton, although p r e v i o u s l y o s s i f i e d i s now mostly cartilaginous. Paddle f i s h they have a l s o Lost most o f t h e i r ganoid scales. Their jaws have degenerated and they tend t o scavenge w i t h t h e i r mouths open. This i s c u r r e n t l y t h e most nunerous group o f vertebrates w i t h respect t o nunber o f species and t o t a l population nunber. From t h e carboniferous time they have become t h e most dominant f i s h group. b These f i s h a r e c h a r a c t e r i z e d by ganoid scales. A ganoid scale i s formed by p a r a l l e l Layers o f bone, above t h i s p o r t i o n was middle Layer o f spongy bone r i c h l y supplied w i t h blood vessels. F i n a l l y there i s an upper Layer o f hard ganoine formed around numerous p u l p c a v i t i e s , above which t h e r e i s a t h i n Layer o f enamel. These are s i m i l a r t o sarcopterygianfs cosmoid scale except t h e L a t t e r have cosmine r e p l a c i n g ganoine, which forms a p a r t i c u l a r l y heavy, shiny scale. B i c h i r (Polypterus) occur i n Central A f r i c a . These f i s h have a symnetrfcal caudal f i n , a dorsal f i n t h a t i s s p l i t i n t o a s e r i e s o f s a i l Like structures, and p a i r e d f i n s t h a t resemble those o f Lobe-fined f i s h . These f i s h a l s o possess Lungs, and t h e nasal s t r u c t u r e i s s i m i l a r t o the coelacanths. The scales a r e a heavy ganoid type. These were o r i g i n a l l y thought t o belong t o t h e Crossopterygians, but recent i n t e r p r e t a t i o n i s t h a t they a r e modified descendants o f the Palaeoniscoids. HOLOSTEI Over e v o l u t i o n a r y time t h e a c t i n o p t e r y g i a n f i s h Lost the various Layers o f t h e scale, w i t h time i t f i n a l l y came t o c o n s i s t s o f o n l y bone. I n p r i m i t i v e a c t i n o p t e r y g i a n f i s h t h e r e were f u n c t i o n a l Lungs (and even i n one L i v i n g f i s h today) but otherwise these have been m o d i f i e d i n t o h y d r o s t a t i c organs (swim bladders) which c o n t r o l t h e buoyancy o f t h e animal). From an e a r l y stage these f i s h had Large eyes, and probably . These were t h e dominant f i s h o f t h e middle Mesozoic. Some had Long, up-turned s h a r k - l i k e t a i l s . The jaws became shorter, and t h e scales tended t o Lose t h e i r ganoid coverings. Most o f t h e r a d i a t i o n was i n t h e marine environment, and w i t h the f o l l o w i n g exceptions t h i s group i s e x t i n c t . Gar pikes (Lepisosteus), these are f a s t swimning f i s h t h a t are s i m i l a r t o the ancestral forms, they have elongated jaws and are associated w i t h a predacious habit. (m), these f i s h are more derived ( v i z Bowfin less p r i m i t i v e than t h e gar pikes), t h e i r scales have l o s t t h e i r ganoid covering. ALL modern Amphibians belong t o the SUBCLASS LISSAMPHIBIA. SALAMANDERS (ORDER CAUDATA) F Head and neck region d i s t i n c t , t h e trunk Long and e i t h e r c y l i n d r i c a l o r depressed dorsoventrally and possess a Long t a i l . TOADS AND FROGS (ORDER ANURA) TELEOSTE I b . Originated i n t h e middle t o l a t e Mesozoic. E a r l y forms s u p e r f i c i a l l y resembled sharks, but t h e i r upturned shark- 1ike heterocercal t a i 1s became reduced and they have a symnetric appearance ( t h e v e r t e b r a l axis i s nevertheless turned upwards). The pectoral f i n s are positioned high up on t h e l a t e r a l surfaces and t h e p e l v i c f i n s are positioned f a r forward. The scales have l o s t a l l o f t h e i r ganoid coverings. The reasons f o r t h e success o f t h e t e l e o s t f i s h are locomotion and feeding mechanisms. During the e v o l u t i o n the jaws became s h o r t e r as the hyomandibular bone and other elements involved i n the suspension o f the jaws r o t a t e d t h e i r p o s i t i o n on the b r a i n case. As t h i s occurred t h e m a x i l l a gradually became f r e e from the other bones and t h e Lever r o t a t e d about i t s a n t e r i o r attachment. The r e s u l t o f t h i s i s t h a t the jaw system became adapted t o almost any type o f d i e t . Teleosts are t h e most v e r s a t i l e group and comprise over 20 000 species. These animals are able t o adapt t o eating micro-organisms r i g h t through t o some h i g h l y predacious forms such as t h e f a s t s w i m i n g tuna f i s h . We even have f i s h t h a t crawl over mud (mudskippers) and f l y out o f t h e water ( f l y i n g f i s h ) . CLASS AMPHIBIA (Frogs, Several features adapt amphibians f o r t e r r e s t r i a l L i f e such as legs, Lungs, n o s t r i l s connecting t o the mouth cavity, and sense organs t h a t can f u n c t i o n i n both water and a i r . The t r a n s i t i o n form water t o Land involves the following:- b b b b CAECILIANS, LEGLESS FORMS (ORDER GYMNOPHIONA) b Limbless and worm l i k e , w i t h a s k i n containing many small i n t e r n a l scales. EXTERNAL FEATURES OF THE ANURAN Highly vascularized s k i n w i t h many mucous glands, t h i s maintains a moist body surface f o r cutaneous respiration. Skin may contain poison glands f o r protection. Protection may be associated w i t h aposematic coloration. Changes i n c o l o r a t i o n can occur due t o three types o f chromatophores. Yellowish hue Iridophores: L i g h t r e f l e c t i n g and produce a l i g h t s c a t t e r i n g (Tyndall Blue) which a c t i n g with yellow pigments produces a green coloration. Melanophores: These are black pigments under the iridophores, and causes t h e s k i n t o darken. When t h e melanin granules move outward they obscure t h e iridophores. toads, sa lamanders .?i caeci 1ians) I n t h e l a t e Palaeozoic amphibians f l o u r i s h e d (Carboniferous t o e a r l y Permian). E a r l y amphibians were put i n t o a subclass Lepospondyli, and were f a i r l y small and s i m i l a r t o modern salamanders. This group even includes forms t h a t have reverted t o l o s i n g t h e i r Limbs. Another group o f amphibians became very Large animals (up t o 3 m i n length and were c a l l e d Labyrinthodonts), and were probably ancestral t o t h e amphibians o f today. The most p r i m i t i v e f o s s i l amphibians c e r t a i n l y belong t o t h e Subclass Labyrinthodontia and include an animal c a l l e d Ichthyostegalia, f o s s i l s o f which were found i n Greenland. These animals possessed a Long f i s h - l i k e t a i l and were very s i m i l a r t o t h e Crossopterygian fish, except they were t h e f i r s t amphibian t o walk on Land. b Head and trunk are joined i n a broad depressed body, with no neck o r t a i l , s h o r t forelimbs, Long h i n d limbs and exposed ear druns. M o d i f i c a t i o n f o r t r a v e l , p l u s maintaining swimning. Change i n s k i n t o f a c i l i t a t e respiration. Replace g i l l s with lungs Changes i n t h e c i r c u l a t i o n system permits r e s p i r a t i o n by lungs and skin. Sense organs t h a t are Land/water compatible. Larvae o f the amphibian such as t h e frogs have m u l t i p l e a o r t i c arches as i n fishes. A f t e r metamorphosis they Lose a l l but one p a i r o f arches. The amphibian s k u l l i s much less complex than t h a t o f t h e f i s h and possess fewer bones. The l i m b muscles become more complex i n . the a d u l t frog. The p i t u i t a r y hormone INTERMEDIN causes dispersal o f the melanin and c o n t r a c t i o n o f t h e iridophores. VERTEBRA Up t o 100 i n t h e anurans 250 i n t h e c a e c i l i a n s PELVIC GIRDLE Well developed i n the anurans Less w e l l developed i n t h e salamanders Without g i r d l e and Limbs i n c a e c i l i a n s Most amphibians have f i n e t e e t h (upper jaw and roof o f mouth) and a long prehensile tongue which i s s i t u a t e d as f a r forward as possible. Segmented muscles are conspicuous on t h e trunk and t a i l i n t h e Salamanders and i n t h e trunk o f t h e Caecilians. G i l l bearing species have special muscles t o move the g i l l s and open o r close t h e g i l l s l i t s . RESPIRATION b Have the most nunber o f techniques f o r r e s p i r a t i o n o f a l l vertebrates. This r e f l e c t s the t r a n s i t i o n from aquatic t o t e r r e s t r i a l conditions. The f o l l o w i n g are used f o r respiration:- g i l l s , skin, Lungs, buccopharynx, and may operate separately o r i n conjunction with each other. w The heart i n f r o g Larvae i s f i s h l i k e w i t h one atrium and a v e n t r i c l e : i t receives only directly to unoxygenated blood which i s pu@ the g i 11s. Adults have two a t r i u m and one v e n t r i c l e . The l e f t a t r i u m receives oxygenated blood from the s k i n and Lungs, t h e r i g h t unoxygenated blood from t h e general c i r c u l a t i o n . Despite t h e s i n g l e v e n t r i c l e there i s some separation o f a t r i a l blood: t h a t sent t o the r e s p i r a t o r y surfaces i s almost a l l unoxygenated. Three p a i r s o f external g i l l s occur i n most embryos and Larvae but p e r s i s t only i n those a d u l t forms t h a t are s t r i c t l y aquatic (u. Sa lamander) I n Larval tadpoles water i s drawn i n through t h e mouth and n o s t r i l s and i s then forced over t h e g i l l s and out through the spiracles. I n salamanders they a i d t h e i r r e s p i r a t i o n by moving t h e i r g i l l s . (u frogs) have Those amphibians w i t h Lungs few i n t e r n a l p a r t i t i o n s t h a t are vascularized. Consequently t h e Lungs are not very e f f i c i e n t . I n aquatic species lungs serve as h y d r o s t a t i c organs being i n f l a t e d when the animals are f l o a t i n g ( t h e r e i s Lung reduction i n some Salamanders). The s k i n o f a l l amphibians i s h i g h l y vascularized and t h i s permits some aquatic species t o remain submerged f o r Long periods and even hibernate i n ponds. b Many species have buccopharyngeal respiration; p u l s a t i o n s o f the t h r o a t move a i r i n and out o f t h e mouth c a v i t y and aeration o f the blood occurs i n vessels i n t h e mucous membrane. REPRODUCTION b Mating u s u a l l y occurs i n t h e water, w i t h external f e r t i l i z a t i o n ( f r o g s and toads). eggs a r e deposited and grow u n t i l they metamorphose i n t o the a d u l t stages. The The male enters t h e water, clasps the female, and as the female extrudes her eggs the male discharges sperm over them. Some Salamanders have elaborate courtship where t h e male deposits a gelatinous spermatophore on t h e bottom o f the stream. The spermatophore i s taken i n t o t h e female cloaca where t h e spermatozoa are stored i n the seminal receptacle and are f e r t i l i z e d ( i n t e r n a l l y ) a t a l a t e r stage. b The amphibian eggs are covered i n a gelatinous coating, w i t h a d i s t i n c t i v e form o f egg mass and j e l l y coating. LARVAE b Salamanders resemble t h e i r parents i n general form. b Larvae o f toads and f r o g s are the f a m i l i a r tadpole which i n i t i a l l y has external g i l l s , which are then replaced by i n t e r n a l g i l l s . Limbs develop a t a Later stage. The mouth i s horny w i t h Labial t e e t h i s sued t o scrape food material. The i n t e s t i n e i s Long and slender. METAMORPHOSIS INVOLVES I' Growth o f a wide mouth, loss o f horny jaws and thedevelopmentofconicalteeth. Loss o f g i l l s , closure o f g i l l s l i t s and development o f lungs. 3) Development o f t h e p r o t r u s i b l e tongue. 4) Emergence o f t h e f o u r Limbs ( p a i r e d appendages). 5, Reduction i n Length o f t h e i n t e s t i n e from Long herbivore type t o short carnivore type. 6) Re-absorption o f t a i l and median f i n s . 7) Biochemical changes include a change i n the excretory products from t o x i c amnonia t o less t o x i c urea. A change i n t h e capacity o f t h e blood t o release oxygen t o t h e tissues, an adaptation r e f l e c t i n g a more a c t i v e L i f e s t y l e . The carnivorous d i e t requires t h a t p r o t e i n d i g e s t i o n be more important (achieved by increased pepsin and t r y p s i n action). Other enzymes are secreted which are responsible f o r the reabsorption o f t h e t a i l . T i p o f the snout has two small v a l v u l a r n o s t r i 1s CLASS REPTILIA These include the f o l l o w i n g L i v i n g ORDERS:Lizards & snakes Turtles & tortoises Crocodiles & a l l i g a t o r s Tuatara (Sphenodon) Eyes are Large and Lateral (upper and Lower eyelids) and a transparent n i c t i t a t i n g membrane ORDER SQUAMATA ORDER CHELONIA ORDER CROCODILIA ORDER RHYNCHOCEPHALIA There are some t e n orders o f r e p t i l e s t h a t are e x t i n c t and they include animals such as t h e dinosaurs and the pterosaurs ( t h e f l y i n g forms). Opening t o the ears i s behind t h e eye and under a moveable f l a p b The vent i s i n t h e form o f a Longitudinal s l i t behind the base o f the h i n d Limbs. BODY COVERING These were t h e f i r s t group o f vertebrates adapted f o r l i f e i n d r y places. Their d r y horny s k i n and scales r e s i s t Loss o f moisture. b Horny scales arranged i n transverse and Lengthwise rows. GENERAL CHARACTERISTICS b Adult crocodiles have an almost ttexoskeletontt o f separate bony dermal p l a t e s (osteoderms) under the dorsal scales i n t h e r e g i o n o f t h e neck t o the t a i l . These are rectangular o r oval and o f t e n p i t t e d and some have a median keel. There are two p a i r s o f epidermal musk glands. 1) Body i s covered w i t h d r y c o r n i f i e d s k i n w i t h scales o r scutes, there are few surface glands. Two p a i r s o f limbs each t y p i c a l l y w i t h f i v e toes ending i n horny claws. They are s u i t e d f o r running, crawling o r climbing. Limbs may a l s o be paddle-shaped as i n marine t u r t l e s and are reduced i n some Lizards, and are absent i n t h e snakes. 3) The skeleton i s completely o s s i f i e d . 4) The heart i s i m p e r f e c t l y four chambered, there are two a t r i a and a p a r t l y d i v i d e d v e n t r i c l e ( t h e v e n t r i c l e s i n crocodiles i s completely divided) ') Respiration i s v i a lungs; pharyngeal and cloaca1 r e s p i r a t i o n occurs i n some aquatic turtles. 6 Twelves p a i r s o f c r a n i a l nerves 7) The body temperature i n Living forms i s variable (poikilothermic) *) F e r t i l i z a t i o n i s i n t e r n a l v i a copulatory organs; t h e eggs are Large with much y o l k and have e i t h e r a Leathery o r Limy s h e l l (may be r e t a i n e d i n body o f some snakes and Lizards u n t i l the young hatch from t h e i r eggs) SKELETON b Massive s k u l l , w i t h pronounced reduction i n the nunber o f bones. b The long Lower jaw a r t i c u l a t e s a t each s i d e o f the p o s t e r i o r margin o f the s k u l l on the quadrat bone. The v e n t r a l surface o f the cranium i s the Long hard p a l a t e above which are the r e s p i r a t o r y passages. The v e r t e b r a l c o l m consists o f 9 10 5 2 +39 b 9, Segmentation i s meroblastic; and eggs have an amnion, chorion, y o l k sac and a l l a n t o i s . The young when they hatch, resemble t h e adults with no metamorphosis. b b b b Dry s k i n Limbs which permit r a p i d running locomotion. Greater separation o f oxygenated and unoxygenated blood. Complete o s s i f i c a t i o n o f the skeleton. Eggs s u i t e d t o development on Land w i t h membrane and s h e l l s t o p r o t e c t the embryo Between the stermm and t h e pubic bones there are seven p a i r s o f V shaped abdominal r i b s . MUSCULAR SYSTEM b Muscles o f the head, neck and Limbs are w e l l d i f f e r e n t i a t e d , though not as c l e a r l y d i f f e r e n t i a t e d as i n mamnals. DIGESTIVE SYSTEM • Buccal cavity, Large mouth, stout, unspecialized teeth ( f o r offence and defence) b F l a t tongue b Oesophagus: slender tube b Stomach: large spherical d i v i d e d i n t o fundus and p y l o r i c p o r t i o n s b Coiled small i n t e s t i n e b Larger rectun b Cloaca and vent b Accessory organs t o t h e stomach are t h e Liver with two Lobes which Lie a n t e r i o r t o the stomach and a pancreas which l i e s i n the f i r s t duodenal Loop. ADVANCES OVER AMPHIBIANS b Cervical vertebrae (short 8 f r e e r i b s ) Thoracic vertebrae (connected t o sternum) Lunbar vertebrae Sacral vertebrae Caudal vertebrae REPTILES ARE NOT AS ADVANCED AS MAMMALS b b b Lack i n s u l a t e d body coverings No i n t e r n a l r e g u l a t i o n o f body temperature True i n t e r n a l development o f embryo does not occur (eggs are simply retained i n the oviduct) EXTERNAL FEATURES: THE CROCODILE b b Head, neck, trunk and t a i l are c l e a r l y discernable. CIRCULATORY SYSTEM Long mouth and c o n i c a l teeth set i n sockets. b Heart comprises a small sinus venosus, two a t r i a and two v e n t r i c l e s . From blood t r a v e l l i n g i n t h e veins i t enters the sinus venosus -r r i g h t v e n t r i c l e + pulmonary a r t e r y + lungs -r pulmonary veins + l e f t atrium -r L e f t v e n t r i c l e + emerges as a p a i r o f a o r t i c arches. ARTERIAL SYSTEM Carotid arteries Subclavian -r -r SOME CHARACTERISTICS OF OTHER LIVING REPTILES SCALES b neck & head forelimbs A o r t i c arch 8 dorsal aorta cavity -r body COLORATION VENOUS SYSTEM b A n t e r i o r vena cava C o l l e c t s from neck, head 8 forelimbs + reproductive organs 8 kidney Single mid dorsal + p o s t e r i o r vena cava Hepatic p o r t a l + digestive tract TURTLES AND TORTOISES RESPIRATORY SYSTEM Paired n o s t r i l s I n t e r n a l nares P a l a t i n e valve, prevents water entering Glottis Larynx ( c a r t i l a g i n o u s w i t h p a i r e d vocal cords) Tubular trachea r e i n f o r c e d w i t h c a r t i l a g e Divides i n t o two short bronchi Lungs spongy w i t h greater p a r t i t i o n i n g + ureter + cloaca NERVOUS SYSTEM/SENSE ORGANS F Two o l f a c t o r y Lobes connected t o t h e large cerebral hemisphere b Two oval o p t i c Lobes F Median pear-shaped cerebellun b Medulla oblongata spread L a t e r a l l y F Narrows i n t o a s p i n a l cord F V e n t r a l l y between t h e bases o f t h e cerebral hemi sphere are t h e o p t i c t racts/opt ic nerves, infundibulum and hypophysis, twelve p a i r s o f c r a n i a l nerves and p a i r e d spinal nerves. Oval s h e l l composed o f p l a t e - l i k e bones w i t h a d e f i n i t e p a t t e r n sutured together with dense connective tissue. Dorsal surface i s r e f e r r e d t o as a carapace and the v e n t r a l surface i s r e f e r r e d t o as t h e plastron. S o f t - s h e l l e d t u r t l e s have a Leathery carapace. The jaws lack teeth and bear c o r n i f i e d b e a k - l i k e structures t o cut/tear/crush vegetation. SPHENODON (TUATARA) b EXCRETORY SYSTEM Two f l a t l o b u l a r kidneys Reptiles can a l s o have some very d i s t i n c t i v e colour patterns. The chromatophores (pigments) resemble those o f t h e amphibians w i t h X (ye1 low), iridophores (bluish/white) and dark rnelanophores. Colour changes are under hormonal c o n t r o l o f intermedin. SOME LIVING REPTILE GROUPS b b Modified i n a v a r i e t y o f ways such as t h e minute scales on t h e f e e t o f geckoes which allow them t o climb smooth surfaces. Marine iguanas have a c r e s t o f Long scales on t h e i r backs. Chameleons have scales formed i n t o long horns and the A u s t r a l i a n thorny d e v i l has scales modified i n t o thorns f o r protection. L i z a r d - l i k e animal but very much more p r i m i t i v e , found o n l y i n New Zealand. Has a median pineal eye and no copulatory organs. LIZARDS b Varied body shapes. Limbs can be long, short, v e s t i g i a l (reduced) o r absent. The t a i l vertebrae i n some species may be incompletely o s s i f i e d and i f the t a i l i s seized by a predator the vertebrae separates a t these breaking p o i n t s and the t a i l i s broken o f f . Lost t a i l p a r t s can be regenerated. SNAKES b SENSES INCLUDE b Taste buds on the tongue b O l f a c t o r y c e l l s i n nose b Eyes (colour v i s i o n ) w i t h Lachrymal glands b Ears short external a u d i t o r y canal, tympanic membrane, middle ear houses one ear bone (stapes) and the inner ear consists o f three semicircular canals REPRODUCTIVE SYSTEM b Paired gonads b Mature male has two round testes, a ductus deferens and a s i n g l e median penis b Mature female has two ovaries, an open funnel o f t h e oviduct running t o the cloaca Characterized by a loss o f appendages and extreme elongation. Vestiges o f the p e l v i c g i r d l e and h i n d Limbs p e r s i s t i n boas, pythons etc. Most snakes, however, lack Limb girdles, sternum, eyelids, external ear openings and a bladder. Snakes are characterized by nunerous vertebrae (200 t o 4001, many slender segmented body muscles connect t h e vertebrae t o vertebrae, the r i b s t o vertebrae and t h e r i b s t o t h e r i b s . Locomotion without Limbs i s achieved by backward pressure o f t h e Lateral Loops o f t h e body against surface i r r e g u l a r i t i e s . Snakes can t r a v e l i n s t r a i g h t l i n e s by h i t c h i n g scales against t h e ground surface. Snakes can ingest whole, Large-sized prey items. This i s achieved through modifications t o the jaw. This m o d i f i c a t i o n s include two mandibles connected w i t h f l e x i b l e connective tissue, loose attachment o f other jaw bones (quadrate) and movement o f the bones o f the p a l a t e and slender backward p o i n t i n g teeth. I n e f f e c t the snake i s able t o d i s l o c a t e i t s Lower jaw t o increase i t s gape size. hopping, running, climbing o r swirmning. The upper segments o f t h e h i n d Limb are muscular, whereas the slender lower legs o r shanks contain tendons, w i t h L i t t l e muscle and sheathed i n c o r n i f i e d scales, as are the f o u r toes which end i n horny claws. CLASS AVES CHARACTERISTICS . b b b b b b b b Body covered with feathers Two p a i r s o f limbs; a n t e r i o r limbs modified f o r f l i g h t , p o s t e r i o r adapted f o r perching, walking o r swimming. Each f o o t i s u s u a l l y equipped w i t h f o u r toes sheathed w i t h c o r n i f i e d skin. Skeleton, t h i n - w a l l e d bones but possessing great s t r u c t u r a l strength, f u l l y ossified, mouth possessing a p r o j e c t i n g beak o r b i l l made from k e r a t i n i z e d t i s s u e w i t h a horny sheath and teeth are absent, s k u l l w i t h one o c c i p i t a l condyle, neck very f l e x i b l e , p e l v i s fused, sternum enlarged and u s u a l l y possessing a median keel, t a i l vertebrae fused and compressed p o s t e r i o r l y . Heart f o u r chambered (two a t r i a and two separate v e n t r i c l e s ) ; o n l y t h e r i g h t a o r t l c arch persists; red blood c e l l s are nucleated, oval and biconvex. Respiration i s by way o f compact Lungs attached t o r i b s and connected t o t h i n - w a l l e d a i r sacs extending between i n t e r n a l organs, voice formed from t h e s y r i n x a t the base o f t h e trachea. No o r d i n a r y bladder, e x c r e t i o n i n semi s o l i d form ( u r i c acid; females u s u a l l y only have the L e f t ovary and oviduct. Twelve p a i r s o f c r a n i a l nerves Body temperature i n t e r n a l l y regulated (homeothermy) F e r t i l i z a t i o n i n t e r n a l , eggs with large yolk, covered w i t h a hard Limy s h e l l and deposited e x t e r n a l l y f o r incubation. C e l l d i v i s i o n i s meroblastic and embryonic membranes (amnion, chorion, y o l k sac and a l l a n t o i s ) present during development w i t h i n t h e egg. Young hatchlings are cared f o r and feed by parents. F b BODY COVERING b b b b b b b b Insulated body coverings. Complete separation o f venous and a r t e r i a l . c i r c u l a t i o n i n the heart. Regulated body temperature. High r a t e o f metabolic a c t i v i t y Ability to f l y Specialized care f o r young. b b body covering not o f h a i r mode o f reproduction very d i f f e r e n t t h e i r a b i l i t y t o f l y (Limited t o bats i n t h e case o f mamnals) D i s t i n c t head, long f l e x i b l e neck, a stout spindle-shaped body o r trunk. b The mouth i s extended as a pointed biLl/beak w i t h a horny covering. On the upper mandible are two s l i t - l i k e n o s t r i l s b The eyes a r e Large and Lateral, each w i t h an upper and Lower eyelid. Beneath there i s a n i c t i t a t i n g membrane, which can be drawn independently across the eyeball from the a n t e r i o r corner. b Below and behind each eye i s an ear opening, hidden under special feathers. The base o f t h i s feather bud sinks i n t o a c i r c u l a r depression: t h e f u t u r e f o l l i c l e . The outermost epidermal c e l l s on the bud become a smooth c o r n i f i e d sheath (periderm): other epidermal c e l l s become arranged i n p a r a l l e l ribs, a Large median one forming the f u t u r e shaft, the others producing barbs. CONTOUR FEATHERS b The external covering and includes t h e enlarged f l i g h t feathers o f the wings and t a i l . b Consists o f a vane, c e n t r a l shaft, a hollow q u i l l attaching t o the f o l l i c l e . Each h a l f o f the vane i s composed o f many narrow p a r a l l e l and c l o s e l y spaced barbs j o i n i n g t h e sides o f t h e shaft. EXTERNAL FEATURES b Begins Like the scale o f a r e p t i l e : v i z as a Local dermal p a p i l l a t h r u s t i n g up t h e o v e r l y i n g epidermis. The c e n t r a l s o f t , dermal p u l p ( o r i g i n a l dermal p a p i l l a ) contains t h e blood vessels and i s wholly n u t r i t i v e , d r y i n g on completion o f growth, so t h a t the feather i s p u r e l y an epidermal structure. The pigments f o r c o l o r a t i o n are deposited i n c e l l s when t h e growth i s completed, t h e sheath breaks and crumbles away o r i s removed by preening. Birds are notably d i f f e r e n t from mamnals i n b D i s t i n c t i v e epidermal structures. They are Lightweight, f l e x i b l e but r e s i l i e n t body coverings and create a i r pockets which insulates t h e body covering, p r o t e c t s the s k i n and creates wings and t a i l f o r support i n flight. THE GROUTH OF A FEATHER b b The s o f t f l e x i b l e s k i n i s Loosely attached t o t h e muscles beneath. I t Lacks glands except f o r the o i l gland above the base o f t h e t a i l . This gland secretes an o i l y substances f o r I1dressingl1the feathers and keep t h e b i 11 from becoming b r i t t l e . The feathers grow from f o l l i c l e s i n the skin. FEATHERS Birds represent s i g n i f i c a n t advances b The short t a i l bears a f a n - l i k e group o f Longt a i l feathers. On the proximal and d i s t a l s i d e o f each barb are numerous smaller, p a r a l l e l barbules, these are provided w i t h minute b a r b i c e l s o r hooklets, serving t o h o l d opposing rows o f barbules Loosely together. DOWN FEATHERS b b The two forelimbs are modified i n t o wings and are attached high-up on the back and have long f l i g h t feathers. The wings are folded i n t o a Z shape formation a t r e s t and are extended i n flight. The h i n d Limbs are generally adapted t o bipedal b Young chicks and other hatchlings are covered p r o v i d i n g excellent with s o f t downy plunage insulation. b Down feathers have short q u i l l s , reduced s h a f t s and a Long f l e x i b l e barb w i t h a short barbules. b Down i s a l s o present beneath t h e contour feathers on ducks and other water b i r d s . - r i g i d support f o r f l i g h t and permits a s l i g h t expansion/contraction f o r r e s p i r a t i o n . FILOPLUMES b Minute, h a i r l i k e feathers, sparsely d i s t r i b u t e d over the body. b Grow i n c l u s t e r s near t h e f o l l i c l e s o f some contour feathers. b Each has a Long t h r e a d l i k e shaft w i t h a few week barbs and barbules a t the t i p . The r i b s are l a t e r a l and t h e sternun i s v e n t r a l w i t h a Large median keel below which t h e pectoral muscles attach. Each thoracic r i b has a d i s t i n c t v e r t e b r a l and s t e r n a l part, t h e two j o i n i n g n e a r l y a t a r i g h t angle. The f i r s t f o u r o r f i v e t h o r a c i c r i b s has a p o s t e r i o r f l a t process overlapping t h e r i b s behind t o strengthen t h e thorax. BRISTLES b The pectoral g i r d l e consists o f a sword-like scapula Lying p a r a l l e l t o the vertebrae and over the r i b s . The coracoid bone occurs between the scapula and the sternun. Some b i r d s have h a i r - l i k e growths t h a t are modified feathers, each w i t h a short q u i l l and a slender s h a f t w i t h a few v e s t i g i a l barbs a t t h e base. These a r e seen about t h e mouths o f some b i r d s such as f l y c a t c h e r s and serve a t a c t i l e function. C l a v i c l e hanging v e n t r a l l y from t h e scapula: the two c l a v i c l e s are fused a t t h e i r v e n t r a l ends t o form the I1Vl1 shaped f u r c u l a (wishbone) attaching t o t h e sternun. FEATHER COLORATION/PATTERNING/MOULT b Feather's colour r e s u l t ' s c h i e f l y from pigment deposition during growth. b Iridescence i s due t o microscopic, t h i n p l a t e s on t h e surface s t r u c t u r e o f feathers. b Feathers o n l y grow i n defined areas c a l l e d feather t r a c t s . b Feather replacement i s c a l l e d moult, and i s u s u a l l y an o r d e r l y gradual process. Large feathers on t h e wing are moulted i n syrrmetrical p a i r s so t h a t f l i g h t i s not hindered. Ducks and Penguins moult a l l o f t h e i r feathers simultaneously, and they are f l i g h t l e s s u n t i l t h e replacements grow in. The scapula, coracoid and c l a v i c l e meet d o r s a l l y t o form a c i r c u l a r canal as a p u l l e y f o r the tendon. Each forelimb i s attached high on t h e dorsal surface: the humerus p i v o t i n g i n t h e glenoid fossa on t h e coracoid. The fore-arm contains t h e radius and t h e ulna as i n other Land vertebrates, but other wing bones are modified t o promote a s t a b l e support f o r the f l i g h t feathers. There are two carpels and three d i g i t s . - Other carpels are fused t o t h e metacarpals t o form the carpometacarpus SKELETON b D e l i c a t e compared t o t h a t o f mamnals. b Many o f t h e bones possess c a v i t i e s t o Lessen t h e i r weight. The skeleton i s modified f o r f l i g h t , f o r bipedal Locomotion, and t h e Laying o f r e l a t i v e l y Large eggs w i t h hard shells. - t h e second d i g i t bears the a l u l a - two the t h i r d d i g i t i s t h e longest (composed o f segments) - Bones o f t h e cranium are separate i n young, krt are fused i n a d u l t s (except f o r t h e nasal f r o n t a l hinge which permits s l i g h t movement o f t h e upper jaw). The p r i n c i p a l f l i g h t feathers (primaries) are supported on d i g i t s three and f o u r and on the carpometacarpus. The braincase i s rounded, Large o r b i t s occur f o r t h e eye. Jaws formed by (premaxi llae, maxi 1l a e and mandibles) p r o j e c t forward o f the bony beak. The secondary f l i g h t feathers are supported on the ulna The t e r t i a r y f l i g h t feathers are supported on the humerus. Lower jaws hinge on the moveable quadrate connecting t o t h e squamosal bone. The s k u l l a r t i c u l a t e s on a s i n g l e o c c i p i t a l condyle with t h e f i r s t neck vertebrae. The p e l v i c g i r d l e i s broad: u n i t e d w i t h a synsacrum, t h a t i s widely opened v e n t r a l l y , p e r m i t t i n g the passage o f large eggs. 16 c e r v i c a l vertebrae (neck) each i s saddle shaped and permits f r e e movement o f the neck f o r feeding and preening. The trunk vertebrae are c l o s e l y f i t t i n g , they have r i b a r t i c u l a t i o n s t h a t are Lateral i n t h e thorax region, but are otherwise fused i n t o a s o l i d synsarcun t o which the p e l v i s i s attached. the f o u r t h d i g i t i s t h e innermost (composed o f a s i n g l e segments) The p e l v i c g i r d l e consists o f an a n t e r i o r iliun, a p o s t e r i o r ischiun, a slender v e n t r a l pubis where these three bones meet, and a socket where the acetabulun receives the head o f the femur. b The Leg consists o f a -- Tibiotarsus Femur - No Lumbar region i s evident. - Four f r e e caudal vertebrae. - Compressed terminal pygostyle (fused) permits movement o f t h e t a i l . SLender/incomplete F i b u l a Fused tarsometatarsus Patella: h e l d i n Ligaments before femurtibiotarsal Four toes: three i n f r o n t and one behind. MUSCULAR SYSTEM A Large t h o r a c i c c a v i t y formed from r i b s which p r o t e c t s t h e i n t e r n a l organs and provides a b Lower vertebrates have dominance o f segmented a r t e r i e s each w i t h three l a r g e branches: the c a r o t i d t o t h e head/neck, t h e branchial t o the wing and t h e pectoral t o t h e breast. The arch continues as the dorsal aorta supplying t h e r e s t o f the body. over non-segmented muscles. B i r d s and mamnals have a greater proportion o f non-segmented muscles. The Large pectoral muscles which o r i g i n a t e on the outer side o f t h e s t e r n a l keel and i n s e r t s on t h e v e n t r o l a t e r a l head o f t h e humerus. A h e p a t i c - p o r t a l system exists. t I n other Land vertebrates the forelimb i s r a i s e d by muscles on the dorsal surface, but i n b i r d s such movement i s due t o a v e n t r a l muscle c a l l e d t h e supracoracoideus. t N o s t r i l s : connected t o i n t e r n a l nares and a slit-like glottis t Trachea: r e i n f o r c e d w i t h h o o p - l i k e c a r t i l a g e that i s p a r t l y calcified. t Syrinx: possess vocal muscles. b Bronchus: 4 lungs which are small interconnecting chambers t h a t opening i n t o Larger chambers c a l l e d the Parabronchi which comnunicate with t h e bronchi and a i r sacs which extend between organs i n t h e body c a v i t y and space around t h e neck vertebrae and i n t o t h e Larger bones. RESPIRATORY SYSTEM - supracoracoideus o r i g i n a t e s on t h e keel and tapers t o a strong tendon. The muscles o f t h e femur and t h e t i b i o t a r s u s are used f o r running and perching. The shank and f e e t c o n t a i n l i t t l e muscle: an adaptation t o reduce heat Loss. Toes are moved by tendons t h a t are connected t o muscles i n the upper segments o f the Legs. Tendons move through spaces Lubricated by fluid. - Air i s brought i n by movement o f muscles between the r i b s bowing L a t e r a l l y and by abdominal muscle. DIGESTIVE SYSTEM - A renal p o r t a l system i s reduced. The c o n t r a c t i o n o f t h e pectoral muscle moves t h e wing downwards and l i f t s the b i r d 1 s body i n flight. - Contraction o f t Tongue, small and pointed with horny covering - w Mouth c a v i t y : roofed w i t h Long p a l a t a l f o l d s t Short pharynx - t Tukrlar/muscular oesophagus t Stomach: s o f t a n t e r i o r proventriculus t Disk shaped v e n t r i c u l u s o r gizzard + walls o f Lined i n t e r n a l l y by t h i c k , dense musculature e p i t h e l i a l s e c r e t i o n -, food ground up by a c t i o n o f muscular w a l l and w i t h gravel/stones g& which f u n c t i o n as teeth. t - - I n t e s t i n e i s slender and consists o f several c o i l s o f the Large rectum a t t h e junction o f the two caeca o r b l i n d pouches. t Cloaca: d o r s a l l y bears an outgrouth: t h e Bursa o f Fabricus. t Large r e d Liver i s b i - l o b e d w i t h a g a l l bladder and two b i l e ducts. t Pancreas: three ducts discharge i n t o t h e a n t e r i o r Loop o f t h e i n t e s t i n e . HEART t - - t h e thorax reverse t h e flow. On i n s p i r a t i o n a i r enters the bronchus, by passes t h e Lungs and enters the p o s t e r i o r a i r sac. On expiration, i t flows i n t o t h e small Lungs. Upon the next i n s p i r a t i o n a i r w i l l f i l l the p o s t e r i o r a i r sacs, t h e f i r s t a i r w i l l pass from the Lungs t o the a n t e r i o r sacs. U i t h e x p i r a t i o n a i r i n t h e a n t e r i o r sac i s expelled i n t o the bronchus and t o t h e outside through the mouth. I n t h i s way a one-way flow o f a i r i s maintained through t h e Lungs. The d i r e c t i o n o f a i r i s opposite t o t h a t o f blood c r e a t i n g a counter-current system. A i r sacs may a l s o d i s s i p a t e heat generated through the muscular contractions. EXCRETORY SYSTEM t Paired kidneys: three-lobed and attached d o r s a l l y under t h e p e l v i s . t From each i s a slender u r e t e r which extends p o s t e r i o r l y t o the dorsal w a l l o f the cloaca. t No bladder t Semi-solid nitrogenous waste ( u r i c acid) pass down the u r e t e r s and out o f the cloaca v i a the vent. NERVOUS SYSTEM AND SENSE ORGANS Four chambered, two t h i n - w a l l e d a t r i a and two t h i c k walled v e n t r i c l e s . Proportionately larger than r e p t i l e s . t O l f a c t o r y lobes are small (poor sense o f smell). t Cerebral hemispheres a r e l a r g e and smooth. Completely separate venous/arterial bloodstreams. Sinus venosus i s incorporated i n t h e r i g h t atriun. Blood from two precaval and one post caval enters t h e r i g h t atrium -, r i g h t v e n t r i c l e -, pulmonary a r t e r y -, c a p i l l a r i e s o f the Lung -, l1oxygenatedl1 r e t u r n s v i a pulmonary veins t o L e f t a t r i u m + L e f t v e n t r i c l e -, s i n g l e r i g h t a o r t i c arch which gives o f f two innominate Optic lobes on t h e midbrain are w e l l developed. t Cerebel lun has an increased surface w i t h many s u p e r f i c i a l folds. t Nerve cord and p a i r e d s p i n a l nerves s i m i l a r t o other vertebrates. - .# b The t h o r a c i c o r branchial plexus serves the Large p e c t o r a l muscles f o r f l i g h t . s h e l l . The s h e l l i s incubated during growth o f the embryo. b- The eyes are p r o p o r t i o n a t e l y very Large: phenomenally keen s i g h t . Colour v i s i o n i s acute and focusing very rapid. The young hatchlings o f chickens and ducks are precocia1:- they are well formed and f u l l y covered w i t h down and are able t o run away idiately. b Hearing i s not as acute as f o r m a m l s . A short external a u d i t o r y canal -, leads t o tympanic membrane -, bone c o l u n e l l a a u r i s which transmits waves across t h e oval window -, From t h e middle ear on each s i d e i s a eustachian tube Leading t o the pharynx with t h e two having a comnon opening a t the back o f t h e palate. -, t h e cochlea o f t h e inner ear i s a short, blind-ended tube t h a t i s intermediate i n the developnent between r e p t i l e s and mamnals. -, Chemoreception (smell and taste) i s u n l i k e l y t o be w e l l developed except i n a few cases. Chicks o f song birds, canaries, pigeons hatch as b l i n d , naked and helpless and are c a l l e d a l t r i c i a l chicks. B i r d s generally have d i f f e r e n t and c h a r a c t e r i s t i c seasons f o r breeding. Breeding comnences w i t h a c o u r t s h i p display, defence o f t e r r i t o r i e s , followed by c o n s t r u c t i o n o f nests, Laying o f eggs, incubation o f eggs and t h e caring o f t h e hatchlings. FLIGHT b The shape o f the wings and t h e manner i n which they are moved r e s u l t s i n forward propulsion w i t h as L i t t l e expenditure o f energy as possible. ENDOCRINE GLAND b P i t u i t a r y : below t h e base o f t h e brain. b Thyroid: two l a t e r a l Lobes beneath the jugular and a t t h e o r i g i n where the subclavian and c a r o t i d meet. b The wing i s an a i r f o i l w i t h a Leading edge t h a t i s t h i c k and a t r a i l i n g edge t h a t i s t h i n . The upper surface o f t h e wing i s curved convexly, whereas t h e Lower surface i s s l i g h t l y concave. Adrenals: v e n t r a l surface o f t h e kidneys Pancreas: w i t h i s l e t s o f Langerhans. n ;. b The spindle-shaped stream-lined body r e s u l t s i n m i n i m a i r resistance. Consequently a i r passing over t h e curved upper surface must move f a s t e r than t h e a i r underneath the wing. Endocrine secretions o f t h e gonad regulate plumage). secondary sexual characters ( s REPRODUCTIVE SYSTEM This causes a s l i g h t negative (Low) pressure on the upper surface i n r e l a t i o n t o t h e lower surface. MALE b Two oval, w h i t i s h t e s t e s a r i s i n g from each i s a much convoluted ductus deferens which extends p o s t e r i o r l y p a r a l l e l t o the ureter. I n many b i r d s i t i s d i l a t e d as seminal vesicles before e n t e r i n g cloacal. Some b i r d s have v e s t i g i a l penis (non-functional). These d i f f e r e n c e s i n pressure creates an upward force c a l l e d l i f t (generated by t h e need t o equalize the pressures on e i t h e r s i d e o f the wing; since the wing has a h i g h pressure on the underside and a low pressure on t h e upper surface, the force generated i s upwards). During breeding t e s t i s becomes enlarged, sperm accumulates i n t h e ductus deferens and i s t r a n s f e r r e d t o t h e cloaca o f the male during copulation. By increasing the angle o f attack ( r a i s i n g the p o s i t i o n o f t h e Leading edge i n r e l a t i o n t o the t r a i l i n g edge) causes an increased amount o f L i f t u n t i l a s t a l l i s induced. FEMALE b System develops o n l y on the Left side, t h e ovary i s ear t h e L e f t kidney: Close by i s t h e open expanded funnel o r infundibulun o f the oviduct. The duct extends p o s t e r i o r l y t o the cloaca. I n non-laying birds, the ovary i s small containing minute eggs and the duct i s small. I n the season o f egg-laying the ovary enlarges, a mature ovum receives a f u l l quota o f y o l k before being released. The mature ovum escapes from t h e ovarian f o l l i c l e i n t o the abdominal c a v i t y and enters t h e funnel a t the top end o f t h e oviduct (movement i s by c i l i a t e d epithelium). F e r t i l i z a t i o n occurs i n the upper oviduct, albumen i s added by glands i n the middle p o r t i o n and t h e s h e l l membranes are secreted by glands i n t h e p o s t e r i o r part. REPRODUCTION (BIRDS IN GENERAL) b A s t a l l i s where t h e smooth a i r flow i s disrupted and turbulence i s induced, the r e s u l t i s t h a t no L i f t now occurs and f l y i n g i s no Longer possible. Eggs are l a i d w i t h much y o l k and a hard Limy U n l i k e a fixed-wing a i r c r a f t , t h e a i r f o i l can be changed i n b i r d s by muscular actions. The outerpart o f t h e major f l i g h t feathers ( t r a i l i n g edge) are s e m i f l e x i b l e and can be separated, c r e a t i n g s l o t s which reduce t h e turbulence and avoid a s t a l l when there i s a high angle o f attack. ( s i m i l a r t o f l a p s i n f i x e d wing a i r c r a f t ) . b The shape o f t h e wing i s r e l a t e d t o t h e type o f f l i g h t . Warm a i r r i s i n g from sun-heated s o i l s o r rocks provides upward thermals i n which b i r d s c i r c l e t o maintain p o s i t i o n on the thermal e.g.vultures. Such b i r d s have broad, large wings. Wind deflected upwards by mountains permits a s i m i l a r soaring p a t t e r n o f f l i g h t . I n the more usual, h o r i z o n t a l winds, soaring b i r d s may remain s t a t i o n a r y o r move forward. B i r d s such as albatrosses soar close over t h e ocean i n an undulating pattern, r i s i n g against ever present winds and they use long narrow wings. Flapping f l i g h t i s t h e most usual. The wings are f u l l y spread and swept forward ( a c t i o n o f t h e p e c t o r a l muscle) and then are p a r t l y folded f o r t h e recovery s t r o k e t h a t i s upwards and backwards. F a i r l y small wing characterize such fliers. SUBCLASS PROTOTHERIA ORDER MONOTREMATA b These are the most p r i m i t i v e , s u r v i v i n g mamnals o f today. b The o n l y l i v i n g Order: MONOTREMATA b Before hatching t h e embryo l i v e s on t h e y o l k contained w i t h i n the p l i a b l e egg. b A f t e r hatching t h e young are nourished by m i l k from the mother's m a m r y glands. CLASS MAMMALIA b b This r e f e r s t o animals possessing mamnary glands. b Mamnals possess body h a i r ( a t sane time i n t h e i r L i f e h i s t o r y , although t h i s may be secondarily Lost) b They are Homeothermic:- maintaining a constant body temperature. b They are endothermic:- meaning t h a t the heat f o r maintaining body temperature comes from t h a t generated w i t h i n the body. b M a j o r i t y o f mamnals a r e viviparous, with t h e young u s u a l l y nourished before b i r t h through a placenta t h a t attaches t h e embryo t o the uterus. * 9 There a r e approximately 4000 L i v i n g species o f mama ls . ' EVOLUTION b Evolved from mamnal-like r e p t i l e s (synapsids). b The synapsid l i n e eventually Lead t o the therapsids, one Line o f which were the theriodonts. The theriodonts were half-way between a r e p t i l e and a mamnal, and were small animals occupying s i m i l a r L i f e - s t y l e s t o t h e modern day shrews. b - The d i s t i n c t i v e mamnalian c h a r a c t e r i s t i c s t h a t evolved w i t h these e a r l y m a m l s included an endothermic physiology w i t h associated homeothermic a t t r i b u t e s such as i n s u l a t i n g hair, increased c i r c u l a t i o n e f f i c i e n c y , more e f f e c t i v e Locomotion, v i v i p a r i t y and greater parental investment i n t h e i r young. b With the e x t i n c t i o n o f the dinosaurs 70 m i l l i o n years ago, many p o t e n t i a l l1nichesl1on Land, sea and i n t h e a i r were vacated and an explosive species r a d i a t i o n o f m a m l s followed during t h e Cenozoic period. b The mamnals are d i v i d e d i n t o two L i v i n g subclasses the P r o t o t h e r i a (egg-laying p r i m i t i v e mamnals) and the Theria (more modern viviparous mamnals). b The t h e r i a group are diverged i n t o the Metatheria (pouched o r marsupial animals) and t h e Eutheria (placental rnamnals). MAMMAL IA - PROTOTHERIA (monotremes) - THERIA -- METATHERIA (marsupial EUTHERIA (placental) There are no teats; the young Lap up m i l k as i t oozes from the tubular glands. w There are f i v e L i v i n g species o f spiny anteater (Echidna) and one species o f d u c k - b i l l e d platypus (Orni thorhynchus). b They Left the main m a m l i a n stock f a r back the Mesozoic era. b The s k u l l has may p r i m i t i v e and/or r e p t i l i a n features. The vertebra are very Like t h a t o f r e p t i l e s , e s p e c i a l l y t h e c e r v i c a l vertebrae which bear separate r i b s as occurred i n the synapsid r e p t i l e s . b The d i f f e r e n t i a t i o n o f t h e atlas, a x i s and seven c e r v i c a l vertebrae i s t h e Least f o r any mamnalian group. b There are 17 and 16 t h o r a c i c vertebrae f o r t h e platypus and the spiny anteater respectively. b Limbs are supported on p e c t o r a l and p e l v i c girdles, and more s i m i l a r t o those o f a r e p t i l e , and have f a r Less muscle than the average advanced mamnal. in Both the platypus and spiny anteater are h i g h l y specialized animals and have no teeth i n t h e i r a d u l t phase, t h e r e f o r e we cannot deduce the primitiveness o f these. The p a t t e r n o f reproduction i s unique amongst mamnals and involves Laying o f eggs. The eggs have undergone p a r t i a l cleavage p r i o r t o being Laid. They show the beginning o f t h e process o f s u b s t i t u t i o n o f post-natal n u t r i t i o n from ovarian n u t r i t i o n . Eggs are incubated f o r 10 days. The female platypus makes long burrows where the nest i s made. At b i r t h the young rnonotremes possess an "egg toothr1 a unique character f o r mamnals. A f t e r incubation and hatching t h e young enter the pouch and are feed by m i l k . Postnatal care obviously preceded t h e egg Laying characteristic. Both the echidna and t h e platypus produce m i l k from about 120 Large s p e c i a l i z e d sweat glands on the v e n t r a l abdomen. The are no ducts u n i t i n g t h i s secretion. ALL other mamnals have nipples which do u n i t e such secretions. Although p r i m i t i v e these animals have h a i r suggesting that mesozoic mamnals and even synapsid mamnals had made progress w i t h temperature regulation. Although p r i m i t i v e these animals are able t o regulate t h e i r body temperatures a t around032 0C i n an ambient temperature range o f 5 t o 30 C. The epipubic bones are reduced i n t h e f u l l y quadrupedal t e r r e s t r i a l forms such as the Tasmanian Wolf. A curious f e a t u r e found i n monotremes i s a grooved e r e c t i l e poison spine on t h e tarsus o f t h e male. The c i r c u l a t i o n system i s r e l a t i v e l y advanced. I n conclusion t h e monotremes are mamnals with respect t o t h e i r brain, h a i r (insulation), warm-bloodedness, heart, Larynx and diaphragm. However, they have obvious r e p t i l i a n a f f i n i t i e s w i t h respect t o t h e i r ovoviviparous reproduction and t h e i r skeleton. b MARSUPIAL REPRODUCTION b The p a t t e r n i s very d i f f e r e n t t o t h a t o f placentals. › Pregnancy i s very short, even less than t h e oestrus cycle. Hormones normally associated w i t h development during pregnancy o f placentals can be removed i n marsupial animals without terminating the pregnancy. SUBCLASS THERIA INFRACLASS METATHERIA ORDER MARSUPIALIA These are t h e pouched animals and are represented by some 250 L i v i n g species. Many o f these species show a strong anatomical resemblance t o placental animals, except f o r t h e i r mode o f reproduction. b b Marsupials diverged from an e a r l y stage i n the e v o l u t i o n o f t h e mamnals. There i s no special imnune suppressant system p r o t e c t i n g the foetus from t h e mothers antibodies as has t o occur i n p l a c e n t a l mamna 1s . b A t h i n s h e l l p r o t e c t s t h e o w n from t h e responses o f the uterus t o the f a t h e r ' s antigens. This p r o t e c t i o n i s b r i e f and t h e young are born w i t h i n 10 t o 12 days o f the s h e l l breaking down. b The L i f e o f t h e mother i s not i n t e r r u p t e d by any dramatic b i r t h . The young g a i n Less from the mother before b i r t h than occurs i n placentals. The marsupial mother r i s k s Less and r e t a i n s more freedom. b A f t e r b i r t h t h e young enjoy Long p r o t e c t i o n i n the pouch. These young can be e a s i l y abandoned i f t h e mother i s stressed. The marsupials p a r a l l e l the adaptive r a d i a t i o n accomplished elsewhere by t h e placental mamnals except f o r forms l i k e bats, whales o r seals. L i k e t h e monotremes many o f t h e i r features are s p e c i a l i z e d so t h a t they represent not a s i n g l e stage on t h e way t o the placental evolution, but a s p e c i a l i z e d s i d e branch. There are 172 marsupial species i n Australia, and accounts f o r t h e e n t i r e mamnalian fauna o f t h i s continent (except f o r aquatic seals and a species o f b a t ) and t h e monotremes. There are successful representatives i n North and South America e.g. opossuns which have survived i n competition w i t h the placentals. Although temperature c o n t r o l may be less well developed than i n the placental animals, Learning s k i l l s , v i s u a l d i s c r i m i n a t i o n and problem s o l v i n g i s equal t o t h a t o f s i m i l a r p l a c e n t a l mamnals. The s k u l l i s s i m i l a r t o t h a t o f the p r i m i t i v e placentals such as the insectivores, with a r a t h e r small craniun. The teeth are not easy t o i n t e r p r e t t h e primitiveness o f the group. The i n c i s o r teeth are more nunerous than i n p l a c e n t a l m a m l s . There are three and four premolar and molar teeth respectively i n each s i d e o f t h e upper o r lower jaw whereas i n p l a c e n t a l there are f o u r and three premolars and molars respectively. Obviously many forms have l o s t teeth and t h i s represents the ancestral condition. The form o f llcuspstt on the teeth do some primitiveness. The skeleton i s generally s i m i l a r t o t h a t o f t h e placentals and has g r e a t l y changed from the monotremes and the r e p t i l e s . The thoracic region consists o f 13 p a i r s o f r i b - b e a r i n g vertebrae as i n placentals and there are u s u a l l y 7 lumbar vertebrae. The pectoral g i r d l e i s s i m i l a r t o t h a t o f the placentals, whereas the p e l v i c g i r d l e d i f f e r s w i t h t h e presence o f the epipubic bones (may be r e f e r r e d t o as the marsupial bones) and i s s i m i l a r t o t h a t o f the monotremes. These epipubic bones are two Large, f l a t bones, which curve forwards and outwards from the m i d l i n e and have the general form o f p a r t i a l crescents. These bones are o n l y l i g h t l y joined t o other bones, and take up the special stresses produced by the abdominal muscles and pouch. The hands u s u a l l y c a r r y f i v e d i g i t s and armed with claws, but t h e nunber o f toes i s o f t e n reduced and may bear h o o f - l i k e structures. The marsupial egg has a very unequal cleavage, there i s v i r t u a l l y no placental development and instead u t e r i n e m i l k may be taken up by the y o l k sac. The embryo may be born as young as 8 days from conception (e.g.the opossuns). At b i r t h the young embryo w i l l leave the vaginal opening and crawl t o t h e pouch and become attached t o the teat. I n order t o make t h i s journey t h e embryo ( c a l l e d a neonate) has well-developed forelimbs and nervous system. I t mouth and o l f a c t o r y system are w e l l developed and i s h i g h l y specialized f o r suckling. I n the pouch the neonate attaches t o t h e teat, and t h e sides o f the Lips grow around the teat, making i t almost impossible t o remove. M i l k i s then i n j e c t e d i n t o the neonate. The Length o f time i n the pouch i s v a r i a b l e between marsupial forms. A wallaby may remain some 8 t o 9 months i n i t s mother's pouch. Marsupials also have delayed implantation. Copulation can take place w i t h i n a day o f b i r t h , but any 80 c e l l b l a s t o c y t e can remain dormant f o r up t o 11 months. suspension o f development i s c o n t r o l l e d by suckling o f the young i n the pouch. Suckling stimulates a release o f oxytocin and there are high l e v e l s o f p r o l a c t i n which i n h i b i t s f u r t h e r development o f the corpus Luteun. PLACENTAL MAMMALS SUBCLASS THERIA INFRACLASS EUTHERIA EIGHTEEN ORDERS - INSECTIVORA - DERMOPTERA MACROSCELIDA SCANDENTIA CHIROPTERA PHOLIDOTA TUBULIDENTATA EDENTATA PRIMATES - - ROOENTIA - - - LAGOMORPHA CETACEA - - - CARNIVORA PROBOSClDEA HYRACOlDEA SIRENlA - - - PERISSODACTYLA ARTIODACTYLA SlRENIA - - small, p r i m i t i v e insect-eating f l y i n g Lemurs elephant shrews t r e e shrews bats phol idota aardvarks edentata Lemurs, t a r s i e r s , monkeys, a p s , hunans gnawing animals except r a b b i t s and hyraxes rabbits whales, dolphins and porpoises modern meat-eating elephants hyraxes seacows, manatees and dugongs odd-toed ungulates even-toed ungulates seacows, manatees and dugongs genetic material from the father. When i t becomes connected t o the mother's body, i t r i s k s imnunological r e j e c t i o n . Nevertheless, placental mamnals can remain i n the uterus, even u n t i l they are capable o f f u l l m o b i l i t y . Even a f t e r b i r t h they are provided w i t h m i l k f o r a f u r t h e r period. b The most widespread and diverse modern marmlals are the placentals, a group whose young complete t h e i r embryonic development attached t o a placenta i n s i d e the mother's uterus. b ORIGINS b < E a r l i e s t forms found i n t h e cretaceous period and were a l l small insect-eating forms. During t h e Palaeocene both carnivore and primates evolved, i n the Late Cretaceous/Palaeocene the ungulates had evolved. b CHARACTERISTICS OF PLACENTAL MAMMALS b A b The placental breeding technique spares the young the hazardous journey outside t h e i r mothers body a t a very e a r l y age. The placenta permits the mother t o supply t h e i r every need during the long g e s t a t i o n period. Animals such as a whale o r seals could not have evolved through the marsupial Lines. Order i n s e c t i v o r a includes are v a r i e t y o f insect-eating forms. There moles which Live i n underground tunnels, hedgehogs which are spiny and r o l l i n t o a b a l l as a defence against predation, the desmans which swim w i t h webbed f e e t and have a long snout which act as a snorkel and shrews which are small a c t i v e predators on insects. I n Madagascar there i s a group c a l l e d the tenrecs, which are extemely diverse being aquatic and t e r r e s t r i a l and some even resemble hedgehogs. Order dermoptera (= skin-winged) evolved from insectivores and are comnonly c a l l e d f l y i n g lemurs. These animals do not have t r u e wings, but a f u r r e d s k i n extending from behind the ears outwards t o the d i g i t s , along the sides o f the body t o the t a i l . These f l a p s o f s k i n only allow them t o g l i d e r a t h e r than t o f l y . Possess n e i t h e r marsupial bones nor a pouch. The foetus develops d i r e c t l y and e n t i r e l y w i t h i n the body o f t h e female. The embryo i s connected i n d i r e c t l y t o the mother v i a a placenta t o t h e w a l l o f a uterus. b A placenta permits t h e embryo ( c a l l e d a foetus) t o remain i n contact w i t h i t r s mothers uterus f o r a Long time. A placenta i s a f l a t d i s k t h a t becomes attached t o the wall o f the uterus and i s connected t o the foetus by an umbilical cord. Order Macroscelidia are a l s o c l o s e l y a l l i e d t o the insectivores and are comnonly c a l l e d the elephant shrews which are h i g h l y specialized f o r running w i t h very Large hindlimbs. Their eyes are b e t t e r developed than t h e poorly sighted insectivores. b Order scandentia include animals c a l l e d t r e e shrews (Tupaia). They are an intermediate group between the i n s e c t i v o r e s and the primates (apes and monkeys). b Order chiroptera includes t h e o n l y t r u e f l y i n g mamnals the bats. Although they have obvious i n s e c t i v o r e origins, t h e i r e n t i r e bone s t r u c t u r e i s modified f o r f l i g h t . The wings are formed by the elongated f i n g e r bones over which i s tretched a naked s k i n membrane. Bats can be dived i n t o small carnivores forms and Large f r u i t eating forms. The small carnivorous bats hunt using echolocation. This bats emit high frequency c l i c k i n g sounds, which s t r i k e objects and the echo i s returned t o the bats using i t s Large ears t o receive them. b Order pholidota are comnonly c a l l e d pangolins o r scaly anteaters. They possess Large scales which are modified h a i r s and s h i e l d the animal form predators. They a l s o have no teeth, but do have an exceptionally Long s t i c k y tongue (anchored t o the p e l v i s ) which i s used t o eat ants and termites. b Order tubulidentata are comnonly c a l l e d aardvarks and specializes i n e a t i n g ants and termites and only occur i n southern A f r i c a . L i k e the pangolins they have Long s t i c k y tongues, but they are not toothless and possess f i v e upper and f o u r lower cheek t e e t h on each side o f t h e i r jaws ( i n c i s o r and canines are The j u n c t i o n o f t h e placenta with the u t e r i n e w a l l i s h i g h l y convoluted so t h a t the surface area between the placenta and maternal t i s s u e I t i s here t h a t the interchange i s v e r y great. between t h e mother and foetus takes place. Blood i t s e l f does not pass from the mother t o i t s foetus, but oxygen from her Lungs and n u t r i e n t s derived from her food are both dissolved i n her blood, d i f f u s e across the j u n c t i o n and so enters i n t o the blood o f the foetus. The waste products produced by the foetus are absorbed by the motherrs blood and then excreted through her kidneys. THE PLACENTAL CONDITION MAKES FOR COMPLICATIONS, ONE OF THE MOST SIGNIFICANT BEING BIOCHEMICAL. The mamnalian sexual c y c l e involves the regular production o f a new egg. This causes no problem t o t h e marsupial since the neonate emerges before the next egg i s due t o be produced. Since t h e p l a c e n t a l remains much Longer i n the uterus, t h e placental foetus promotes the s e c r e t i o n o f hormones which suppresses the motherrs sexual c y c l e during pregnancy. The f o e t u s r s tissues are not the same g e n e t i c a l l y as t h e mother's since i t contains absent). the e a r l y Cenozoic. Order edentata are a v a r i e d group t h a t are confined t o t h e new world( North and South America) and includes anteaters, s l o t h s and armadilos. Anteaters are toothless and have s i m i l a r adaptations t o pangolins and aardvarks f o r e a t i n g ants and termites (convergent evolution). Sloths and armadillos lack i n c i s s o r s and canines, but do have s i n p l e cheek t e e t h t h a t Lacks enamel and grow continuously from t h e i r roots. The armadillo has 100 such teeth. The a r m a d i l l o has bony p l a t e s on i t s back f o r p r o t e c t i o n against predators. Sloths are slow moving f u r r e d animals t h a t Live up i n t h e trees. The p r e h i s t o r i c gound s l o t h s reach imnense sizes up t o 5 metres (Megatherim) as d i d some armadillos (Glyptodon) whose bony p l a t e s formed a carapace t h a t was about 1,6 m i n length. Order Carnivora are m a t e a t i n g placentals t h a t have prominent canine teeth. These are Long, curved, pointed, very strong and p a r t i c u l a r l y e f f e c t i v e f o r stabbing. Most carnivores are predominantly meat-eaters and a l s o have short, but sharp i n c i s o r teeth f o r nipping. The f o u r t h upper premolar and f i r s t Lower molar teeth have become s p e c i a l i z e d i n t o shearing blades c a l l e d carnassial teeth (these are absent i n aquatic carnivores such as seals). Cats, dogs, Lions Leopards, cheetahs, Hyaenas mongooses, bears, pandas (secondarily adapted t o a herbivorous d i e t ) , seals, walruses, sealions and elephant seals are a l l members o f Carnivora. Order Proboscidea i s represented by two species o f elephant. Elephants are characterized by an imnense size, and a long trunk t h a t i s r e a l l y a h i g h l y modified nose. The tusks are elongated upper incisors. Elephants have no canines o r premolars and t h e i r molars are used t o g r i n d up herbaceous food and are replaced successively from behind as they wear down. They are naked skinned since they l i v e i n t r o p i c a l climates and t h e i r Large ears a c t as r a d i a t o r s t o d i s s i p a t e excess heat. Their e x t i n c t r e l a t i v e s , t h e mamnoths, had f u r and had colonized cold, northern l a t i t u d e s u n t i l they were hunted t o e x t i n c t i o n by p r e h i s t o r i c man. Order primates includes Lemus, monkeys, apes and man. They are e s s e n t i a l l y t r o p i c a l i n d i s t r i b u t i o n and evolved from ancient, arboral i n s e c t i v o r e s sometime i n t h e l a t e Cretaceous period. Their t r e e - d w e l l i n g existence demanded dextrous 1imbs, hands and f i n g e r s ( i n c l u d i n g an opposable thumb), close set eyes w i t h overlapping f i e l d s o f v i s i o n f o r depth perception, e x c e l l e n t eye-hand co-ordination and extended parental care f o r t h e i r young. They are characterized by considerable development o f the brain, a process t h a t eventually r e s u l t e d i n the e v o l u t i o n o f man. The e a r l i e s t ape-man was Australopithecus africanus, a small-brained formed, followed i n sequence by t h e Larger brained Homo h a b i l i s (who used t o o l s ) and Homo erectus (had use o f t o o l s and f i r e ) . From t h i s stock two forms o f modern man Homo sapiens evolved, one c a l l e d Neanderthal man s u r v i v e i n c o l d northern climates during t h e i c e ages, the other form survives today as Homo sapien sapiens. Order rodentia, i s t h e most abundant o f p l a c e n t a l mamnal forms and includes, rats, mice, mole-rats, s q u i r r e l s ( i n c l u d i n g forms t h a t can g l i d e Like t h e f l y i n g lemurs), porcupines, p r a i r i e dogs. They are a l l characterized by a long p a i r o f prominent upper and Lower incisors. These teeth are kept sharp through d i f f e r n t i a l wear o f t h e i r enamel and are v e r y e f f e c t i v e f o r gnawing. Canine teeth are absent. Order Hyracoidea includes t h e hyrax o r dassie, which although s u p e r f i c i a l l y resemble rodents and lagomorphs, are a c t u a l l y more c l o s e l y r e l a t e d t o elephants. They are herbivores with continuously growing chisel-shaped incisors. They f e e t are completely d i s t i n c t i v e , there being f o u r toes on the f r o n t feet, three on the hind, terminating i n small, h o o f l i k e n a i l s . Order Perissodactyla are odd-toed, hoofed animals (=ungulates) and includes, horses, asses, zebras, t a p i r s ( a l l having a s i n g l e d i g i t ) and rhinoceoses (having three d i g i t s ) . They are a l l Large herbivorous animals, and many have elongated Limbs f o r f a s t e r running speeds (horses and zebras). Their cheek teeth are massive, strong and modified f o r g r i n d i n g p l a n t material. b Order A r t i o d a c t y l a are t h e even-toed hoofed animals and include antelope, deer, hippos, g i r a f f e s , camels, Llamas, c a t t l e , sheep, pigs and goats. These are generally herbivorous, and with t h e exceptions o f p i g s and hippos are ruminant animals (having a Large fore-stomach c a l l e d a runen i n which p l a n t f i b r e s are i n i t i a l l y digested). Many are grazing animals and occur i n enormous herds and a l s o have elongated Legs f o r f a s t e r running and t o avoid predation. Their cheek teeth are large, corrplex and designed f o r g r i n d i n g p l a n t material (upper canine and i n c i s o r s are reduced o r absent. b Order S i r e n i a are t h e aquatic dugongs and mantees (sometimes c a l l e d sea cows). They are Large, h a i r l e s s herbivores (possessing o n l y functional cheek teeth) t h a t have t h e i r f o r e l i n ~ b smodified i n t o paddles and no h i n d limbs. Possible share a s i m i l a r ancestry w i t h elephants. Order Lagomorpha includes r a b b i t s and hares and a l l a r e s t r i c t l y ground dwelling forms. Despite t h e i r s i m i l a r i t i e s t o rodents they are not c l o s e l y related. L i k e rodents they do have continuously growing i n c i s s o r s with a t o t a l o f f o u r i n the upper jaw (two o f which are reduced) as opposed t o only two i n the rodents. Order Cetacea includes t h e most aquatic forms o f mamnals; t h e whales, dolphins and porpoises. The blue-whale i s t h e heaviest animal ever t o have l i v e on t h e earth. Their modifications f o r an aquatic existence include Loss o f the hind-limbs, forelimbs have developed i n t o f l i p p e r s , t h e t a i l i s f l a t t e n e d and most have a dorsal f i n , t h e nasal opening (blow hole) i s on t h e t o p o f t h e head, no external ears, h a i r i s almost absent and i n s u l a t i o n i s by way o f blubber. They can be d i v i d e d i n t o toothed forms which are a c t i v e predators, and toothless forms which have Long p l a t e s o f s t i f f h a i r - l i k e material t h a t forms a baleen which acts as a sieve t o f i l t r e plankton. Cetaceans are i n t e l l i g e n t , s o c i a l animals, that can comnunicate w i t h each other and use echolocation t o navigate the waters they swim i n . They probably evolved from insectivores i n