Chordata The Chordata include animals (including you) that have (even if only as an embryo) a notochord and a hollow dorsal nerve cord. The notochord acts as a flexible stiffening rod in animals that retain it as adults (e.g hagfish and lancelets). In adult humans the notochord forms the intervertebral disks. Hagfish Vertebrata During the Cambrian period approx 500 million years ago chordates evolved a head with a skull becoming Craniates and later a more complex nervous system and a vertebral column. Humans, obviously are both craniates and vertebrates. Vertebrata Next major step in evolution of vertebrates was the evolution of jaws about 470 million years ago. Jaws derived from skeletal rods involved in supporting the gills. Jaws an important evolutionary development that opened huge number of opportunities. Huge number of aquatic jawed fishes evolved including ancestors of sharks and the bony fish. 34 .13 Origin of tetrapods The movement of vertebrates onto the land was another big step. First amphibians evolved from a group of bony fish called the lobe-finned fish. Excellent fossils (mainly from Greenland) document the transition. Limbs appear to have first evolved to enable organism to move along the bottom and only later proved useful on land. 34.19 Acanthostega (345 mya fossil from Greenland) early tetrapod The origin of tetrapods and movement onto land There are a suite of challenges associated with terrestrial living that the first tetrapods has to overcome. These included the fact that water supports a fish’s body but air does not, so stronger skeletal structures were needed to support the body and limbs to allow movement. In addition, gills collapse out of water so an alternative breathing system (i.e. lungs) was needed. The origin of tetrapods and movement onto land In addition, there was a constant threat of water loss from the skin or through eggs, which limited the first tetrapods the amphibians to a close connection with water until hard shelled eggs were evolved. The origin of tetrapods and movement onto land The tetrapods (a monophyletic group that includes the amphibians, reptiles, birds and mammals) are all descended from an ancestral lobe-finned fish. Tetrapod limb The tetrapods are defined by their limbs, which have a characteristic structure. Taking the forelimb, for example, there is first a single bone (humerus), then a pair of bones (radius and ulna), next a series of wrist bones (carpals) and finally a set of digits (phalanges). Tetrapod limb The bones found in modern tetrapod limbs are homologous to those in lobe-finned fishes such as Eusthenopteron and Panderichthys of the late Devonian period about 380 mya and to the earliest known tetrapods amphibians such as Acanthostega and Icthyostega (both approx. 360-365 mya) . Figure 25.01a 17.1a Eusthenopteron Ichthyostega Ichthyostega was discovered in the 1930’s in Greenland. Ichthyostega’s legs probably wouldn’t have supported it well on land, but would have enabled it to move around on the bottom in shallow water. It has 7 digits rather than the five of modern tetrapods. It has a strong spine and ribcage, but the hind legs would not have supported it out of water. Acanthostega A contemporary of Ichthyopstega more recently discovered is Acanthostega. It has 8 digits and a very fish like shoulder structure and no wrist. It would not have been able to come out of water and it possessed gills. Like Ichthyostega it probably made its way along the bottom and crawled over vegetation rather than crawling on land. Figure 25.01b 17.1 B and C The origin of tetrapods and movement onto land The transition from lobe-finned fishes fishes to Acanthostega and Ichthyostega has recently been neatly bridged by the discovery of a classic intermediate form Tiktaalik roseae from Greenland. Tiktaalik roseae Discovered in 2004 in Greenland Tiktaalik roseae is an extremely important fossil link in the origin of tetrapods. Described as a “fishapod” Tiktaalik has a mixture of fish and tetrapod characteristics. Tiktaalik roseae Like a fish it has gills and scales. Intermediate characteristics include halftetrapod/half fish limb bones. There is a wrist, but there are fins instead of toes. The inner ear is also intermediate in structure between fish and tetrapods. Like a tetrapod it has lungs, tetrapod rib bones and a mobile neck. Tiktaalik roseae 375 mya Amphibians and reptiles Lobe-finned fishes gave rise to amphibians which in turn gave rise to amniotes which evolved the hard-shelled (amniotic egg) liberating them from need to lay eggs in water as amphibians do. Also evolved less permeable skin. Modern amphibians: Class Amphibia Amphibians are characterized by a moist permeable skin and this limits the environments in which they typically can live as they are constrained by the threat of water loss. Typically they inhabit wet or damp habitats where the humidity is high (e.g., ponds. forest floors in leaf litter). Class Amphibia The moist skin is used extensively for gas exchange and some species have lost their lungs over evolutionary time and depend exclusively on gas exchange across the skin and oral cavity. All amphibians are ectothermic. Class Amphibia The word amphibian means “two lives” and is a reference to the fact that frogs go through metamorphosis from a tadpole stage. In most amphibians, fertilization is external and the male fertilizes the soft eggs as the female sheds them from her body. Class Amphibia Amphibian eggs do not have a hard shell and dry out quickly if not kept in a moist environment. Many species lay their eggs directly in water, but some frogs show more parental care and brood eggs in (depending on the species) the mouth, stomach or on their back. Figure 25.04 Class Amphibia Eggs in frogs develop into tadpoles that have a fishlike tail and external gills. As the tadpole develops, often very quickly in a race against time to escape a pool before it dries up, limbs develop, the tail shortens by reabsorption and the gills are lost as the tadpole metamorphoses into a miniature frog. Figure 25.26 Class Amphibia The amphibians are represented by about 4800 species and include the familiar frogs, toads and salamanders and the less well-known caecilians, which are legless, burrowing amphibians. Amniote origins and classification The possession of a shelled egg unites the mammals, birds and reptiles into a monophyletic group the amniotes. The shelled egg freed the amniotes from the need to reproduce in water that hampered the amphibians ability to spread into harsh environments. The Amniotic egg The amniotic egg is hard shelled and is called an amniotic egg because the embryo develops within a sac called the amnion. The embryo feeds on yolk from a yolk sac and deposits its waste into another sac called the allantois. The allantois and another membrane the chorion together lie against the shell and being richly supplied with blood exchange gases with the outside through the pores in the shell. Figure 26.04 Amniote origins and classification There is considerable disagreement between cladistic and traditional classification of the amniotes. Traditional classification recognizes three classes: Reptilia: reptiles Aves: birds Mammalia: mammals Amniote origins and classification Because the class Reptilia does not include all the descendents of their most recent common ancestor (i.e. the birds are excluded) the reptiles are a paraphyletic group. Birds and crocodilians share a most recent common ancestor and thus form a monophyletic group (the Archosauria), which includes the extinct dinosaurs, but neither is more closely related than the other to the members of the Reptilia Figure 26.02 18.2 Amniote origins and classification Traditional classification considers birds because of their endothermy and feathers to be members of a different grade to the crocodilians and reptiles and so places them in their own class the Aves. Cladistic classification in contrast groups the amniotes on the basis of common ancestry. Amniote origins and classification One of the major characteristics used to classify the amniotes is the structure of the skull and the number of holes it contains. The stem group of amniotes diverged into three lineages in the Carboniferous period (approximately 350 mya). These were the synapsids, anapsids and the diapsids. Figure 28.01 20.1 Figure 26.02 18.2 Figure 26.01 18.1 Modern reptiles The modern reptiles being a paraphyletic group include anapsids and diaspids. The anapsid representatives are the turtles (order Testudines). Most of the remaining reptiles are diapsids and members of the order Squamata which includes the lizards and snakes. Differences between reptiles and amphibians Reptilian skin is dry and scaly, which limits water loss. The reptiles’ amniotic egg frees reptiles from the need to lay eggs in water. Thus they can occupy much drier habitats. Reptilian jaws are more powerful and can apply a crushing grip. Order Squamata: Suborder Sauria the lizards Lizards are a very diverse group that includes terrestrial, burrowing, aquatic, arboreal and even aerial members. Lizards have good vision and an external ear, which snakes lack. They also have eyelids, also a trait that snakes lack. Order Squamata: Suborder Sauria the lizards Most lizards have four limbs, although some species are completely legless. Well known species include: chameleons, geckos, iguanas and monitor lizards, which include the largest species, the Komodo dragon. Figure 26.12 Gecko (note the flattened pads on the toes. Ridges on these pads enable the gecko to cling to smooth surfaces). Figure 26.14 Chameleon catching an insect with its sticky extensible tongue. Order Squamata: Suborder Sauria the lizards Lizards have invaded many of the world’s hottest areas by evolving a suite of adaptations that make survival in deserts possible. These include a thick skin that contains lipids, which reduce water loss, and the excretion of uric acid which minimizes water loss. Reptiles Reptiles are ectothermic and adjust their body temperature by moving from one microclimate to another to bask or cool down. Cold climates do not suit reptiles as there are too few opportunities to warm up. Because they spend relatively little energy keeping warm, ectotherms in general do well in low productivity ecosystems such as tropical deserts and grasslands. Snakes Snakes are limbless and usually lack both the pectoral and pelvic girdles. They have numerous vertebrae, which are shorter and wider than those in other vertebrates and allow them to make undulatory movements. Snakes Snakes are an extremely successful group of predators. Although most have poor vision (with the exception of arboreal species) and limited hearing ability they use other sense organs to track prey. Snakes have pit-like Jacobson’s organs in the roof of the mouth, which are olfactory organs. The forked tongue when extended samples the air and picks up molecules that are delivered to the Jacobson’s organ when the tongue is withdrawn. Snakes Crotaline vipers (pit vipers such as rattlesnakes) have heat-sensitive pit organs on their heads between the nostrils and eyes. These are very sensitive to radiant heat and can detect temperature differences as slight as 0.003ºC. The vipers use the organ to track prey and to aim their strike when biting. Figure 26.24 18.22 Predation Snakes use one of three methods to catch and kill prey. Most catch prey by grabbing it and swallowing it alive. Most such species are quick and concentrate on small easy to handle prey. The other two group kill their prey either by constriction or with venom. Constrictors A variety of snakes including pythons and boas kill by constriction. They coil around their prey and every time the prey breathes out they tighten their coils a little more until the prey can no longer breathe and suffocates. Most constrictors are large, slow-moving ambush predators and the largest snakes, the Anacondas and pythons are all constrictors. Venomous snakes About 20% of all snakes are venomous (although in Australia 80% of snakes are venomous). About 50,00060,000 people die annually worldwide from snake bite, most of them in the Indian subcontinent. Snakes with venom lethal to humans include the vipers (including the American pit vipers) which have large movable tubular fangs at the front of the mouth; elapids (cobras, mambas, coral snakes kraits which have shorter, but permanently erect fangs in the front of the mouth; sea snakes (family Hydrophiidae). Figure 26.25 18.20 Venomous snakes Snake venoms are highly modified salivas and complex in constitution including a variety of proteins and enzymes. Elapid venom is neurotoxic and works by shutting down the respiratory system whereas viper venom is more painful and attacks the vascular system bringing about coagulation of blood and clotting of arteries. Crocodiles and Alligators: Order Crocodilia Modern crocodiles and birds are the only survivors of the Archosaurian lineage that included the dinosaurs. Crocodiles have changed little in almost 200 million years a testament to the success of their design. Crocodiles Crocodiles have their teeth set in sockets a trait found otherwise only in mammals and fossil birds and also like mammals have a complete palate which enables them to breathe even if the mouth is filled with water or food. They are ambush predators that kill by grabbing and drowning their prey. The largest Nile and Estuarine crocodiles (called “salties” in Australia) can exceed 1000 kgs in weight and can attack and kill almost anything. Figure 26.27a 18.24 Modern Birds The other remaining Archosaurs are birds given their own class Aves by traditional taxonomists. There are approximately 8,600 species of birds ranging is size from the tiny bee hummingbird to the ostrich. Characteristics of the birds Feathers are a unique character of birds among living animals, but also found in dinosaurs. Endothermic Skeleton modified for flight. Bones hollow, forelimbs support the wing, ribs with uncinate processes, beak but no teeth, reduced tail. Breathing by lungs and associated air sacs Internal fertilization and hard-shelled amniotic egg Evolution of birds Birds evolved from a group of theropod dinosaurs in the Jurassic period. The oldest known bird fossil is Archaeopteryx lithographica which has a mix of “reptilian” and avian features. Reptilian: long tail, teeth, long clawed fingers Avian: feathers, ribs with uncinate processes, avian shoulder girdle. Archaeopteryx (oldest known fossil bird) Jurassic 150mya Feathers Feathers are what enable birds to fly, but originally are believed to have evolved as a thermoregulatory device. Feathers are lightweight, but strong. The surface of the feather is made up of tightly spaced, overlapping filaments that hook together. Overlapping feathers form the wings with which birds fly. Figure 27.05e 19.4 Structures and adaptations for flight To generate the power to fly birds have evolved very large flight muscles (the supracoracoidueus and the pectoralis), which attach to an enlarged keeled sternum In addition birds have strengthened the skeleton by fusing bones together and reduced their weight in several ways by: Evolving light, hollow bones Reducing the tail Evolving a toothless bill Dispensing with a bladder and excreting uric acid. Figure 27.08 19.6 How birds fly A bird’s wing is an airfoil and is cambered with a slightly convex upper surface and concave under surface. Because air must travel further over the upper surface of the wing than below it must travel faster and thus exerts less pressure above the wing than it does below. The increased pressure below generates lift, the force which keeps the bird up. Figure 27.16 19.13 Lift and thrust In order to fly both horizontal thrust and vertical lift are required. Thrust is mainly generated by the primary feathers (the long ones at the end of the bird’s hand), which on the downstroke twist and acting like a propeller push the air backwards. Lift is mainly generated by the secondary feathers (the inner portion of the wing), which form an airfoil. Early mammals About 300 million years ago amniotes split into two lineages one, the synapsids, ultimately produced the mammals and the other lineage produced the “reptiles,” dinosaurs, and birds. First mammals arose about 200 million years ago. Small and inconspicuous. Dinosaurs dominated Earth until mass extinction 65 million years ago caused by an asteroid impact wiped them out and mammals diversified. Mammals Mammals are characterized by a suite of shared derived characters including Milk Hair Differentiated teeth Single jaw bone (dentary) Middle ear with three bones: incus, malleus, stapes Muscular diaphragm Two occipital condyles (bones at base of skull that articulate with vertebrae). Endothermic (maintain constant high body temperature; evolved independently by birds) Four chambered heart (also evolved independently by birds) Figure 28.02 20.2 Modern mammals After the Cretaceous extinction mammals radiated to occupy niches previously occupied by the dinosaurs. Today there are 26 orders of mammals that can be divided into three groups on the basis of differences in reproduction: The Monotremes: one order Marsupials: seven orders Placental mammals: eighteen orders Monotremes The monotremes are a small order of four species the duck-billed platypus and three species of spiny anteater or echidna found in Australia and New Guinea. They diverged from the line leading to the other mammals in the Jurassic period about 180 mya. Monotremes have several “reptilian” traits Monotreme means single hole and refers to the fact that the urinary tract and reproductive tract empty into a single opening the cloaca just as is the case in lizards and birds. Monotremes In addition, the monotremes lay eggs with a tough leathery shell and possess an interclavicle bone, one found in reptiles, but not in other mammals. However, they clearly are mammals possessing a single dentary, three ear bones, hair, and milk although they lack nipples and the milk seeps from pores over a relatively wide area. Are endothermic, but have a lower metabolic rate than marsupials and eutherians. 34.33 Echidna or Spiny anteater Marsupials There are seven orders of marsupials with just under 300 species that include such animals as possums, kangaroos, wombats, koala and Tasmanian devil. All marsupials have an abdominal pouch in which the young are raised having been born very underdeveloped, moving to the pouch and latching onto a nipple. Figure 28.24 20.18 34.34 Marsupials Most marsupials are found in Australia (isolated from eutherians for about 65 million years) and the remaining species (opossums) occur in the Americas. Rock Wallaby Convergent evolution of eutherian and marsupial mammals 34.35 Eutherian mammals The Eutherian or placental mammals support their developing young using a chorioallantoic placenta which brings the blood supplies of mother and offspring into close contact so that food and gases can be effectively exchanged. Major orders of eutherian mammals There are about 5000 species of placental mammal that belong to 18 orders. Major orders of mammals include: Insectivora: shrews, hedgehogs, moles. Small, primitive insect eating mammals with many pointed teeth. 440 species Figure 28.16 20.28 Major orders of eutherian mammals Chiroptera: Bats. Flying mammals with forelimbs modified into wings. Membrane stretched between elongated fingers. Many species use echolocation. 977 species second only in size to Rodentia. Rodentia: Rodents. The largest group with 2052 species. Possess chisel like upper and lower incisors. Includes rats, mice, squirrels, woodchucks. Major orders of eutherian mammals Carnivora: large flesh eating mammals. Cats, dogs, weasels, seals, sealions. Teeth specialized for hunting. Canines used for killing. 280 species. Cetacea: whales, dolphins. Aquatic diving mammals. Anterior limbs modified into flippers, posterior limbs absent, possess large rear fluke for propulsion, nostrils modified into blowhole on top of head. 78 species. Figure 28.42 Humpback Whale Figure 28.40 Figure 28.39 Sealion Grizzly Bear Major orders of eutherian mammals Artiodactyla: even-toed ungulates. Pigs, deer, cattle, antelopes, hippopotamuses. Two or four toes sheathed in hoofs. Many are ruminants and possess multi-chambered stomachs. 221 species. Perissodactyla: odd-toed ungulates. Horses, zebras, tapirs, rhinoceroses. One or three toes. All herbivorous with teeth adapted to chewing vegetation. 17 species. Figure 28.41 20.32 Major orders of eutherian mammals Primates: the order to which humans belong includes prosimians, monkeys, apes and us. Largely arboreal with binocular vision and grasping hands. 279 species. Figure 28.31 Tarsier Red howler monkey Figure 28.32b Olive baboon 20.22 20.23 Figure 28.32a Humans are primates The order Primates split from other mammals about 55 mya. Humans are apes and we are most closely related to chimpanzees. Human and chimp lineages separated about 7 million years ago. Primates have grasping hands with a thumb (monkeys apes and humans have an opposable thumb) and binocular vision. These are adaptations to a tree-dwelling existence. Human derived characteristics Humans are bipedal and have an extremely well developed brain. Humans have reduced jawbones and a short digestive tract Genomes of humans and chimps about 99% identical.