GLY1101: Study Guide Nr. 3

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1 Phanerozoic Eonothem: This Eonothem is often referred to as the time of "Visible Life". Organisms with skeletons or hard shells appeared by the first time in the geological record. The Phanerozoic Eonothem spans from 543 mya through today. The Phanerozoic is divided into three Erathems, from older to younger: Paleozoic, Mesozoic, and Cenozoic. The Mesozoic Erathem: 248 to 65 mya The Mesozoic is divided into three time systems: the Triassic ( Million Years Ago), the Jurassic ( Million Years Ago), and the Cretaceous ( Million Years Ago). Mesozoic means "middle life", and is the time during which the world fauna changed drastically from that which had been seen in the Paleozoic. Dinosaurs, which are perhaps the most popular organisms of the Mesozoic, evolved in the Triassic, but were not very diverse until the Jurassic. Except for birds, dinosaurs became extinct at the end of the Cretaceous. Some of the last dinosaurs to have lived are found in the late Cretaceous deposits of Montana in the United States. The Mesozoic was also a time of great change in the terrestrial vegetation. The Lower Mesozoic was dominated by ferns, cycads, ginkgophytes, bennettitaleans, and other unusual plants. Modern gymnosperms, such as conifers, first appeared in their current recognizable forms in the early Triassic. By the middle part of the Cretaceous System, the earliest angiosperms had appeared and began to diversify, largely taking over from the other plant groups. The breakup of Pangaea can be divided into four stages: a. The first stage involved the separation of North America from Africa during the Late Triassic, followed by the separation of North America from South America. b. The second stage involved the separation of Antarctica, India, and Australia from South America and Africa during the Jurassic. During this stage, India broke away from the still united Antarctica and Australia landmass. c. During the third stage, South America separated from Africa, while Europe and Africa began converging. d. In the last stage, Greenland separated from North America and Europe. The Lower Mesozoic = Triassic + Jurassic Sytems The Lower Mesozoic is an informal division of the Mesozoic Erathem which includes the lower two systems: the Triassic and the Jurassic. See Chronostratigraphic Chart Paleogeography: Triassic < Jurassic < Cretaceous < Summary of Lower Mesozoic Events: Important facts in the history of Mesozoic life The Mesozoic is divided into three time systems: the Triassic ( Million Years Ago), the Jurassic ( Million Years Ago), and the Cretaceous ( Million Years Ago). Mesozoic means "middle life", and is the time during which the world fauna changed drastically from that which had been seen in the Paleozoic. Dinosaurs, which are perhaps the most popular organisms of the Mesozoic, evolved in the Triassic, but were not very diverse until the Jurassic. Except for birds, dinosaurs became extinct at the end of the Cretaceous. Some of the last dinosaurs to have lived are found in the late Cretaceous deposits of Montana in the United States. The Mesozoic was also a time of great change in the terrestrial vegetation. The Lower Mesozoic was dominated by ferns, cycads, ginkgophytes, bennettitaleans, and other unusual plants. Modern gymnosperms, such as conifers, first appeared in their current recognizable forms in the early Triassic. By the middle part of the Cretaceous System, the earliest angiosperms had appeared and began to diversify, largely taking over from the other plant groups. 1

2 The breakup of Pangaea can be divided into four stages: a. The first stage involved the separation of North America from Africa during the Late Triassic, followed by the separation of North America from South America. b. The second stage involved the separation of Antarctica, India, and Australia from South America and Africa during the Jurassic. During this stage, India broke away from the still united Antarctica and Australia landmass. c. During the third stage, South America separated from Africa, while Europe and Africa began converging. d. In the last stage, Greenland separated from North America and Europe. The breakup of Pangaea influenced global climatic and atmospheric circulation patterns. While the temperature gradient from the tropics to the poles gradually increased during the Mesozoic, overall global temperatures remained equable. Among the marine invertebrates, survivors of the Permian extinction diversified and gave rise to increasingly complex Mesozoic marine invertebrate communities. Triassic and Jurassic land plant communities were composed of seedless plants and gymnosperms. Angiosperms, or flowering plants, evolved during the Early Cretaceous, diversified rapidly, and soon became the dominant land plants. Dinosaurs evolved during the Upper Triassic, but were most abundant and diverse during the Jurassic and Cretaceous. Based on pelvic structure, two distinct orders of dinosaurs are recognized Saurischia (lizard hipped) and Ornithischia (birdhipped). Pterosaurs were the first flying vertebrate animals. Small pterosaurs were probably active, wing flapping fliers, while large ones may have depended more on thermal updrafts and soaring to stay aloft. At least one pterosaur species had hair or feathers, so it was likely endothermic. The fish eating, porpoise like ichthyosaurs were thoroughly adapted to an aquatic life. Female ichthyosaurs probably retained eggs within their bodies and gave birth to live young. Plesiosaurs were heavy bodied marine reptiles that probably came ashore to lay eggs. Crocodiles become the dominant freshwater predators during the Jurassic. Turtles and lizards were present during most of the Mesozoic, but by the Cretaceous snakes evolved from lizards. Birds probably evolved from small carnivorous dinosaurs. The oldest known bird, Archaeopteryx, appeared during the Jurassic, but few other Mesozoic birds are known. Protoavis in Triassic rocks may represent a bird older than Archaeopteryx. The earliest mammals evolved during the Upper Triassic, but they are difficult to distinguish from advanced cynodonts. Details of the teeth, the middle ear, and lower jaw are used to distinguish the two. Several types of Mesozoic mammals existed, but all were small, and their diversity was low. A group of Mesozoic mammals known as eupantotheres gave rise to both marsupials and placentals during the Cretaceous. Because the continents were close together during much of the Mesozoic and climates were mild even at high latitudes, animals and plants dispersed widely. Mesozoic mass extinctions account for the disappearance of dinosaurs, several other groups of reptiles, and a number of marine invertebrates. One hypothesis holds that the extinctions were caused by the impact of a large meteorite. Many paleontologists reject the meteorite proposal and claim that withdrawal of epeiric seas and climatic changes can account for these extinctions. Introduction to the Tetrapoda: The Four Legged Vertebrates The word "Tetrapoda" means "four legs" in Greek. Amphibians, reptiles (including dinosaurs and birds) and mammals are the major groups of the Tetrapoda. The Dinosauria Dinosaurs, one of the most successful groups of animals (in terms of longevity) that have ever lived, evolved into many diverse sizes and shapes, with many equally diverse modes of living. The term "Dinosauria" was invented by Sir Richard Owen in 1842 to describe these "fearfully great reptiles," specifically Megalosaurus, Iguanodon, and Hylaeosaurus, the only three dinosaurs known at the time. The creatures that we normally think 2

3 of as dinosaurs lived during the Mesozoic Erathem, from late in the Triassic period (about 225 million years ago) until the end of the Cretaceous (about 65 million years ago). But we now know that they actually live on today as the birds. Some things to keep in mind about dinosaurs: Not everything big and dead is a dinosaur. All too often, books written (or movies made) for a popular audience include animals such as mammoths, mastodons, pterosaurs, plesiosaurs, ichthyosaurs, and the sail backed Dimetrodon. Dinosaurs are a specific subgroup of the archosaurs, a group that also includes crocodiles, pterosaurs, and birds. Although pterosaurs are closely related to dinosaurs, they are not true dinosaurs. Even more distantly related to dinosaurs are the marine reptiles, which include the plesiosaurs and ichthyosaurs. Mammoths and mastodons are mammals and did not appear until many millions of years after the close of the Cretaceous period. Dimetrodon is neither a reptile nor a mammal, but a basal synapsid, i.e., an early relative of the ancestors of mammals. Not all dinosaurs lived at the same time. Different dinosaurs lived at different times. Despite the portrayals in movies like King Kong and Jurassic Park, no Stegosaurus ever saw a Tyrannosaurus, because Tyrannosaurus didn't appear on the scene until 80 or so million years following the extinction of stegosaurs. The same goes for Apatosaurus ("Brontosaurus") it was already well fossilized by the time T. rex came along. The Dinosaurs are not extinct: Technically, based on features of the skeleton, most people studying dinosaurs consider birds to be dinosaurs. This shocking realization makes even the smallest hummingbird a legitimate dinosaur. So rather than refer to "dinosaurs" and birds as discrete, separate groups, it is best to refer to the traditional, extinct animals as "nonavian dinosaurs" and birds as well as birds, or "avian dinosaurs." It is incorrect to say that dinosaurs are extinct, because they have left living descendants in the form of cockatoos, cassowaries, and their pals just like modern vertebrates are still vertebrates even though their Cambrian ancestors are long extinct. So at thanksgiving what you eat is a dinosaur we called a turkey The Evolutionary Process of Phanerozoic Life Major groups of animals are already present in the Phanerozoic, at the beginning of the Cambrian System, thier phylogenetic development is shown in the following diagrams as shown below: The opisthokonts (Greek:?????? (opisth? ) = "rear, posterior" +?????? (kontos) = "pole" i.e. flagellum) are a broad group of eukaryotes, including both the animal and fungus kingdoms, together with the phylum Choanozoa of the protist kingdom. Both genetic and ultrastructural studies strongly support that opisthokonts form a monophyletic group. One common characteristic is that flagellate cells, such as most animal sperm and chytrid spores, propel themselves with a single posterior flagellum. This gives the groups its name. In contrast, flagellate cells in other eukaryote groups propel themselves with one or more anterior flagella. See Biological Principles for a review of these concepts Anapsida: amniote group whose skull does not have openings near the temples. The clade Anapsida includes turtles and all their extinct relatives. The anapsids once consisted of many groups, many of which could be considered to have been quite successful until their extinction. Today, only one group of anapsids remains ( Chelonia the turtles), which truly could be called an evolutionary success story. Diapsids ("two arches"): are a group of tetrapod animals that developed two holes (temporal fenestra) in each side of their skulls, about 300 million years ago during the late Carboniferous period. Living diapsids are extremely diverse, and include all birds, crocodiles, lizards, snakes, and tuataras. All members of the group called the Reptilia, except for the anapsids (turtles and their ilk), and a few extinct groups, are diapsids. The main diagnostic physical character for a diapsid is the presence of two openings on each side of the skull; the upper and lower temporal openings. Even the birds are considered diapsids (and hence reptiles), because they are descended from certain dinosaurs (which are also diapsids), and ancestrally have the paired skull openings along with other 3

4 physical characteristics that unite them with diapsids. Thus, they are considered diapsids by their ancestry, which is illuminated by shared derived traits. Synapsids ('fused arch') also known as Theropsids ('beast face'), and traditionally described as 'mammal like reptiles', are a group of amniotes (the other being the sauropsids) that developed one opening in their skull (temporal fenestra) behind each eye, they appeared about 320 million years ago (mya) during the late Carboniferous Period. The mammals of today are but one branch of the Synapsida, a great vertebrate group with a 300 million year history. Pre mammalian synapsids including the famous "finback" Dimetrodon dominated the land vertebrate fauna of the Permian and early Triassic before losing ground to the diversifying dinosaurs and other archosaurs. These pre mammalian groups of synapsids have at times been called "mammal like reptiles". This term is now discouraged because although many had characteristics in common with mammals, none of them were actually reptiles. Euryapsida are a group of tetrapod animals that are distinguished by a single opening behind the orbit (temporal fenestra). They are different from Synapsida by the precise placement of the opening below. It is now commonly believed that euryapsids are in fact diapsids (having two fenestrae) that lost the upper temporal fenestra % of marine species became extinct: The K/T Extinction Paleogeography: Cretaceous < The K/T < Organisms affected: dinosaurs, ammonites, planktonic foraminifera Causes of the K/T Extinction: Plate Tectonics: Climatic Fluctuations: Volcanic Eruptions: Asteroid Impact: The Origin of Vertebrates A chordate (phylum Chordata) is an animal that has, at least during part of its life cycle, a notochord, a dorsal hollow nerve cord, and gill slits. Vertebrates, which are animals with backbones, are simply a subphylum of chordates. The ancestors and early members of the phylum Chordata were soft bodied organisms that left few fossils. As a result, we know little about the early evolutionary history of the chordates or vertebrates. Surprisingly, a close relationship exists between echinoderms and chordates, and they may even have shared a common ancestor (see phylogeny tree above). This is because in the developing embryo of echinoderms and chordates, cells divide by radial cleavage so that the cells are aligned directly above each other. In all other invertebrates, cells undergo spiral cleavage, which results in having cells nested between each other in successive rows. The Fishes: The most primitive vertebrates are fish, and some of the oldest fish remains are found in the Upper Cambrian Deadwood Formation in northeastern Wyoming. Here phosphatic scales and plates of Anatolepis, a primitive member of the class Agnatha (jawless fish), have been recovered from marine sediments. All known Cambrian and Ordovician fossil fish have been found in shallow, nearshore marine deposits, whereas the earliest nonmarine (freshwater) fish remains have been found in Silurian strata. This does not prove that fish originated in the oceans, but it does lend strong support to the idea. As a group, fish range from the late Cambrian to the present. 4

5 The Agnatha: are the oldest and most primitive of class of fish and typified by the ostracoderms, whose name means «bony skin" are. These are armored, jawless fish that first evolved during the late Cambrian, reached their zenith during the Silurian and Devonian, and then became extinct. The majority of ostracoderms lived on the seafloor. A typical examples of ostracoderm are the genera Hemicyclaspis and Pteraspis Hemicyclaspis is a bottom dwelling ostracoderm. Vertical scales allowed Hemicyclaspis to wiggle sideways, propelling itself along the seafloor, and the eyes on the top of its head allowed it to see such predators as cephalopods and jawed fish approaching from above. While moving along the sea bottom, it probably sucked up small bits of food and sediments through its jawless mouth. Pteraspis, was more elongated and probably an activeswimmer, although it also seemingly fed on small pieces of food that it was able to suck up. Primitive jawed fish: The evolution of jaws was a major evolutionary advance among primitive vertebrates. Although their jawless ancestors could only feed on detritus, jawed fish could chew food and become active predators, thus opening many new ecologic niches. The vertebrate jaw is an excellent example of evolutionary opportunism. Various studies suggest that the jaw originally evolved from the first three gill arches of jawless fish. Because the gills are soft, they are supported by gill arches of bone or cartilage. The evolution of the jaw may thus have been related to respiration rather than to feeding. By evolving joints in the forward gill arches, jawless fish could open their mouths wider. Every time a fish opened and closed its mouth, it would pump more water past the gills, thereby increasing the oxygen intake. The modification from rigid to hinged forward gill arches let fish increase both their food consumption and oxygen intake, and the evolution of the jaw as a feeding structure rapidly followed. The fossil remains of the first jawed fish are found in Lower Silurian rocks and belong to the acanthodians (Class Acanthodii), a group of small, enigmatic fish characterized by large spines, paired fins, scales covering much of the body, jaws, teeth, and greatly reduced body armor. Although their relationship to other fish is not well established, many scientists think the acanthodians included the probable ancestors of the present day bony and cartilaginous fish groups. The acanthodians were most abundant during the Devonian, declined in importance through the Carboniferous, and became extinct during the Permian. The other jawed fish, the placoderms (Class Placodermii), whose name means "plate skinned," evolved during the Late Silurian. Placoderms were heavily armored, jawed fish that lived in both freshwater and the ocean, and, like the acanthodians, reached their peak of abundance and diversity during the Devonian. The placoderms showed considerable variety, including small bottom dwellers, as well as large major predators such as Dunkleosteus, a Late Devonian fish that lived in the midcontinental North American epeiric seas. It was by far the largest fish of the time, reaching a length of more than 12 m. It had a heavily armored head and shoulder region, a huge jaw lined with razor sharp bony teeth, and a flexible tail, all features consistent with its status as a ferocious predator. Ages of Fish: Besides the abundant acanthodians, placoderms, and ostracoderms, other fish groups, such as the cartilaginous and bony fish, also evolved during the Devonian. For this reason the Devonian is informally called the "Age of Fish," because all major fish groups were present during this time period. The cartilaginous fish: (Class Chrondrichthyes), represented today by sharks, rays, and skates, first evolved during the early Devonian, and by the late Devonian, primitive marine sharks such as Cladoselache were quite abundant. Cartilaginous fish have never been as numerous or as diverse as their cousins, the bony fish, but they were, and still are, important members of the marine vertebrate fauna. The bony fish: (Class Osteichthyes) also first evolved during the Devonian. Because bony fish are the most varied and numerous of all the fishes, and because the amphibians evolved from them, their evolutionary history is particularly important. There are two groups of bony fish: the common ray finned fish (Subclass Actinopterygii) and the less familiar lobe finned fish (Subclass Sarcopterygii). The term ray finned refers to the way the fins are supported by thin bones that spread away from the body. From a modest freshwater beginning during the Devonian, ray finned fish, which include most of the familiar fish such as trout, bass, perch, salmon, and tuna, 5

6 rapidly diversified to dominate the Mesozoic and Cenozoic seas. Present day lobe finned fish are characterized by muscular fins. The fins do not have radiating bones but rather have articulating bones with the fin attached to the body by a fleshy shaft. Such an arrangement allows for a powerful stroke of the fin, making the fish an effective swimmer. Three orders of lobe finned fish are recognized: coelacanths, lungfish, and crossopterygians. Coelacanths: (Order Coelacanthimorpha) are marine lobe finned fish that evolved during the middle Devonian and were thought to have gone extinct at the end of the Cretaceous. In 1938, however, a fisherman caught a coelacanth in the deep waters off Madagascar and since then, several dozen more have been caught, both there and in Indonesia. Lungfish: (Order Dipnoi) were fairly abundant during the Devonian, but today only three freshwater genera exist, one each in South America, Africa, and Australia. Their present day distribution presumably reflects the Mesozoic breakup of Gondwana. The «lung" of a modern day lungfish is actually a modified swim bladder that most fish use for buoyancy in swimming. In lungfish, this structure absorbs oxygen, allowing them to breath air when the lakes or streams in which they live become stagnant and dry up. During such times, they burrow into the sediment to prevent dehydration and breath through their swim bladder until the stream begins flowing or the lake they were living in fills with water. When they are back in the water, lungfish then rely on gill respiration. The crossopterygians: (Order Crossopterygii) are an important group of lobe finned fish, because it is probably from them that amphibians evolved. However, the transition between crossopterygians and true amphibians is not as simple as it was once portrayed. The group of crossopterygians that appears to be ancestral to amphibians are rhipidistians. These fish, reaching lengths of over 2 m, were the dominant freshwater predators during the late Paleozoic. Eusthenopteron, a good example of a rhipidistian crossopterygian and the classic example of the transitional form between fish and amphibians, had an elongated body that helped it move swiftly through the water and paired, muscular fins that many scientists thought could be used for moving on land. The structural similarity between crossopterygian fish and the earliest amphibians is striking and one of the most widely cited examples of a transition from one major group to another. However, recent discoveries of older lobe finned fish and tetrapods like Acanthostega, and newly published findings of tetrapod like fish, are filling in the gaps in the time of the evolution between fish and tetrapods. Before discussing this transition and the evolution of amphibians, it is useful to place the evolutionary history of Paleozoic fish in the larger context of Paleozoic evolutionary events. Certainly, the evolution and diversification of jawed fish as well as eurypterids and ammonoids had a profound effect on the marine ecosystem. Previously defenseless organisms either evolved defensive mechanisms or suffered great losses, possibly even extinction. Ostracoderms, although armored, would also have been easy prey for the swifter jawed fishes. Ostracoderms became extinct by the end of the Devonian, a time that coincides with the rapid evolution of jawed fish. Placoderms, like acanthodians, greatly decreased in abundance after the Devonian and became extinct by the end of the Paleozoic. In contrast, cartilaginous and ray finned bony fish expanded during the Late Paleozoic, as did the ammonoid cephalopods, the other major predators of the late Paleozoic seas. Amphibians Vertebrates Invade the land Although amphibians were the first vertebrates to live on land, they were not the first land living organisms. Land plants, which probably evolved from green algae, first evolved during the Ordovician. Furthermore, insects, millipedes, spiders, and even snails invaded the land before amphibians. Fossil evidence indicates that such landdwelling arthropods as scorpions and flightless insects had evolved by at least the Devonian. The transition from water to land required animals to surmount several barriers. The most critical were desiccation, reproduction, the effects of gravity, and the extraction of oxygen from the atmosphere by lungs rather than from water by gills. Up until the 1990s, the traditional evolutionary sequence had a Rhipidistian crossopterygian, like Eusthenopteron, evolving into a primitive amphibian like Ichthyostega. At that time, fossils of those two genera were about all paleontologists had to work with, and although there were gaps in morphology, the link between crossopterygians and these earliest amphibians was easy to see. Crossopterygians already had a backbone and limbs that could be used for walking and lungs that could extract oxygen. The oldest amphibian fossils, on the other hand, found in the 6

7 Upper Devonian Old Red Sandstone of eastern Greenland and belonging to such genera as Ichthyostega, had streamlined bodies, long tails, and fins along their backs, in addition to four legs, a strong backbone, a ribcage, and pelvic and pectoral girdles, all of which were structural adaptations for walking on land. These earliest amphibians thus appear to have inherited many characteristics from the crossopterygians with little modification. However, with the discovery of such fossils as Acanthostega and others like it, the transition between fish and amphibians involves a number of new genera that are intermediary between the two groups. Panderichthys, a large (up to 1.3 m long), Late Devonian (~380 million years ago) lobe finned fish from Latvia, was essentially a contemporary of Eusthenopteron. It had a large tetrapod like head with a pointed snout, dorsally located eyes, and modifications to that part of the skull related to the ear region. From paleoenvironmental evidence, Panderichthys lived in shallow tidal flats or estuaries, using its lobe fins to maneuver around in the shallow waters. Acanthostega, a Late Devonian (365 million years ago) tetrapod seemed to be the perfect intermediary between fish and true landdwelling tetrapods. However, its limbs could not support its weight on land, and thus it was an aquatic animal, using its limbs to navigate in water, rather than walking on land. In 2006, an exciting discovery of a m long, 375 million year old (late Devonian) "fishapod" was announced. Discovered on Ellesmere Island, Canada, Tiktaalik roseae, from the Inuktitut meaning "large fish in a stream," was hailed as an intermediary between the lobe finned fish like Panderichthys and the earliest tetrapod, Acanthostega. Tiktaalik roseae is truly a "fishapod" in that it has a mixture of both fish and tetrapod characteristics. For example, it has gills and fish scales but also a broad skull, eyes on top of its head, a flexible neck and large rib cage that could support its body on land or in shallow water, and lungs, all of which are tetrapod features. What really excited scientists, however, was that Tiktaalik roseae has the beginnings of a true tetrapod forelimb, complete with functional wrist bones and five digits, as well as a modified ear region. Sedimentological evidence suggests Tiktaalik roseae lived in a shallow water habitat associated with Late Devonian floodplains of Laurasia. The oldest known amphibian, Ichthyostega, had skeletal features that allowed it to spend its life on land. Because amphibians did not evolve until the Late Devonian, they were a minor element of the Devonian terrestrial ecosystem. Like other groups that moved into new and previously unoccupied niches, amphibians underwent rapid adaptive radiation and became abundant during the Carboniferous and Early Permian. The late Paleozoic amphibians did not at all resemble the familiar frogs, toads, newts, and salamanders that make up the modern amphibian fauna. Rather, they displayed a broad spectrum of sizes, shapes, and modes of life. One group of amphibians were the labyrinthodonts, so named for the labyrinthine wrinkling and folding of the chewing surface of their teeth. Most labyrinthodonts were large animals, as much as 2 m in length. These typically sluggish creatures lived in swamps and streams, eating fish, vegetation, insects, and other small amphibians. Labyrinthodonts were abundant during the Carboniferous when swampy conditions were widespread but soon declined in abundance during the Permian, perhaps in response to changing climatic conditions. Only a few species survived into the Triassic. Evolution of Reptiles: the Land is Conquered Early Reptiles Amphibians were limited in colonizing the land because they had to return to water to lay their gelatinous eggs. The evolution of the amniote egg freed reptiles from this constraint. In such an egg, the developing embryo is surrounded by a liquid filled sac called the amnion and provided with both a yolk, or food sac, and an allantois, or waste sac. In this way the emerging reptile is in essence a miniature adult, bypassing the need for a larval stage in the water. The evolution of the amniote egg allowed vertebrates to colonize all parts of the land, because they no longer had to return to the water as part of their reproductive cycle. Many of the differences between amphibians and reptiles are physiologic and are not preserved in the fossil record. Nevertheless, amphibians and reptiles differ sufficiently in skull structure, jawbones, ear location, and limb and vertebral construction to suggest that reptiles evolved from labyrinthodont ancestors by the Late Mississippian. This assessment is based on the discovery of a well preserved fossil skeleton of the oldest known reptile, Westlothiana, and other fossil reptile skeletons from Late Mississippian aged rocks in Scotland. Other early reptile fossils occur in the Lower Pennsylvanian Joggins Formation in Nova Scotia, Canada. Here remains of Hylonomus are found in the sediments filling in tree trunks. 7

8 These earliest reptiles from Scotland and Canada were small and agile and fed largely on grubs and insects. They are loosely grouped together as protorothyrids, whose members include the earliest known reptiles. During the Permian Period, reptiles diversified and began displacing many amphibians. The reptiles succeeded partly because of their advanced method of reproduction and their more advanced jaws and teeth, as well as their ability to move rapidly on land. The pelycosaurs, or finback reptiles, evolved from the protorothyrids during the Pennsylvanian and were the dominant reptile group by the early Permian. They evolved into a diverse assemblage of herbivores, exemplified by the herbivore Edaphosaurus and carnivores such as Dimetrodon. An interesting feature of the pelycosaurs is their sail. It was formed by vertebral spines that, in life, were covered with skin. The sail has been variously explained as a type of sexual display, a means of protection, and a display to look more ferocious. The current consensus seems to be that the sail served as some type of thermoregulatory device, raising the reptile's temperature by catching the sun's rays or cooling it by facing the wind. Because pelycosaurs are considered the group from which therapsids evolved, it is interesting that they may have had some sort of bodytemperature control. The pelycosaurs became extinct during the Permian and were succeeded by the therapsids, mammal like reptiles that evolved from the carnivorous pelycosaur lineage and rapidly diversified into herbivorous and carnivorous lineages. Therapsids were small to medium sized animals that displayed the beginnings of many mammalian features: fewer bones in the skull, because many of the small skull bones were fused; enlarged lower jawbone; differentiation of teeth for various functions such as nipping, tearing, and chewing food; and more vertically placed legs for greater flexibility, as opposed to the way the legs sprawled out to the side in primitive reptiles. In addition, many paleontologists think therapsids were endothermic, or warm blooded, enabling them to maintain a constant internal body temperature. This characteristic would have let them expand into a variety of habitats, and indeed, the Permian rocks in which their fossil remains are found are distributed not only in low latitudes but in middle and high latitudes as well. As the Paleozoic Erathem came to an end, the therapsids constituted about 90% of the known reptile genera and occupied a wide range of ecologic niches. The mass extinctions that decimated the marine fauna at the close of the Paleozoic had an equally great effect on the terrestrial population. By the end of the Permian, about 90% of all marine invertebrate species were extinct, compared with more than two thirds of all amphibians and reptiles. Plants, in contrast, apparently did not experience as great a turnover as animals. Cephalopods (meaning "head foot") Are mollusks with tentacles and a large head. These soft bodied invertebrates include animals like squid, octopuses, cuttlefish, and the ammonites (extinct). They are fast moving carnivores that catch prey with their tentacles and poison it with a bite from beak like jaws. They move with by squirting water through a siphon, a type of jet propulsion. Many also squirt ink to help escape predators. 8

9 Ammonites were similar in shape and form to the modern Nautilus. They were very common in the oceans of the Paleozoic and Mesozoic erathems, 400 to 65 million years ago, evolving in the Devonian Period. Fossil shells as small as 3 mm and as large as 3 m have been found. Classification Phyllum Mollusca Class Cephalopoda Subclass Ammonoidea Order Goniatida Order Cerartida Order Ammonitida Ammonites are, of course, members of the animal kingdom. Since they are without backbones, they are invertebrates. They belong to the Phylum Mollusca because of their soft body and ability to create shells. Modern day varieties include snails, clams, and oysters, but the octopus and squid have a small internal shell or no shell at all. They are mollusks that belong to the Class Cephalopoda as does the ammonite. The tentacled head is the primary feature of the members of this class, so the modern chambered nautilus also belongs to this class. For our ancient fossil, the subclass is Ammonoidea. These are the orders: Goniatitida The goniatites ranged from the Devonian to the upper Permian. They are characterized by septa with round saddles and pointed lobes. Ceratida This order existed from the Carboniferous to the Triassic. They are characterized by septa with round saddles and serrated lobes. Ammonitida They first show up during the Permian and lasted through the Cretaceous. Their septa have folded saddles and lobes. The shells are decorated with a complex pattern of lines called lirae. The name comes from its appearance: it resembles a rams horn. In Egyptian mythology, the God Ammon looked like a man with horns like a ram. The ancient fossil was considered Ammon s stone, thus inheriting the name, ammonite. Ammonites first appeared in the lower Devonian Period. It is thought by some that they evolved from the older nautiloids. Septa The septa are the walls that divide the chambers within the shell. Nautiloids had simple septa with a single arc. The ammonites developed septa that had intricate folds called lobes and saddles. They also developed delicate lacey patterns on the outer shell. There are three basic patterns for ammonite septa. irregular zigzags this is called goniatite regular wavy called ceratite intricate feathery or fern like patterns ammonite 9

10 The pattern of the septa can be reflected on the outside of the shell. These are called sutures. These patterns along with the shape of the shell and the structure of the septa are how this cephalopod is classified. The geologic distribution of the ammonites GONIATITES (Devonian Permian) had round saddles and mainly undivided pointed lobes. CERATITES (Carboniferous to Triassic) had round saddles and serrated lobes. AMMONITES (Permian to Cretaceous) had folded saddles and folded lobes that are finely subdivided in fractal patterns. 10

11 Chronostratigraphic division of the Mesozoic Cenozoic Erathem Phanerozoic Eonothem Mesozoic Erathem Paleozoic Erathem Triassic = Three fold division of the system in Germany Jurassic = Jura Mountaun, Swiss Alps Cretaceous = Form the latin name for limestone Upper Mesozoic Lower Mesozoic Cretaceous Jurassic Triassic Summary of Mesozoic bioevents Cretaceous Jurassic Triassic Invertebrates: Extinction of ammonites, rudists, and most planktonic foraminifera (Protista) at the end of the Cretaceous Ammonites and belemnoids continue to diversify Rudist become major reef-builders Invertebrates: Ammonites and belemnoid cephalopods increase in diversity Rudist bivalves appear Invertebrates: Survivors of the Permian extinction repopulate the sea Bivalves and echinoids expand into infaunal niche Vertebrates: Extinction of dinosaurs, flying reptiles, and marine reptiles Placental and marsupial mammals diverge Vertebrates: First birds Giant sauropod dinosaurs Vertebrates: Mammals evolved from cynodonts Cynodonts become extinct Ancestral archosaur gives rise to dinosaurs Flying reptiles and marine reptiles evolved Plants: Angiosperms evolve and diversify rapidly Seedless plants and gymnosperms still common but less varied and abundant Plants: Seedless vascular plants and gymnosperms Plants: Land flora of seedless vascular plants and gymnosperms as in Upper Paleozoic Geology: Further fragmentation of Pangaea South America and Africa have separated Australia separated from South American but remains connected to Antarctica North American continues to open Geology: Fragmentation of Pangaea continues, close connections exist among all continents Geology: Fragmentation of Pangaea begins in the upper Triassic 11

12 Reptiles: Early Consumers on Land Evolutionary Novelty: New morphological innovations that define newly established groups: The amniotic egg: a space capsule for the reptilian embryo. The major evolutionary step to a fully terrestrial existence was accomplished primarily due to innovation in reproduction. As one of their diagnostic features, reptiles have an amniote egg, a reproductive character that eventually allowed them to dominate many available land habits. Diagnostic features that developed in the gradual evolution of amphibians to reptiles: Fairly common intermediate forms that combined a blend of typical amphibian and reptilian characters (see figure below). In general, early reptiles stabilized a particular style of backbone construction. The lack of an otic notch in reptiles, the ear being situated at the rear of the skull. Bones of the back part of the reptilian skull are reduced in number and size, a continuation of the trend present from rhipidistian fish to amphibians. A mounted skeleton of the Permian amphibian SeymouriafromCentral Texas (see image). This small animal, approximately 60 cm (2 ft) long, shows a unique blend of amphibian and reptilian characters but is much too young to have been the direct ancestor of reptiles. Notice that an otic notch is still present in the back of the skull. Other anatomical features resemble those of reptiles.(courtesy of NationalMuseum of Natural History.) How to tell early reptiles from others The most important feature for distinguishing one type of early reptile from another is the structure of the bones in the temple region of the skull, behind the eye, called temporal openings, whichprovide data that is used in subdividing all major reptile groups. The reptiles of this group are referred to as stem reptiles or anapsidsbecause they are the ones from which the other, more advanced reptiles are thought to have evolved. The only living anapsids are the turtles and tortoises (see figure below) The synapsids: or mammal like reptiles have a single temporal opening low on the side of the skull, beneath the postorbital and squamosal bones. They are extinct but very important because mammals evolved from this group of reptiles. The diapsids: or ruling reptiles have two openings, one above the other, separated by a bony connection between the postorbital and squamosal bones. They include the dinosaurs of the Mesozoic Era, as well as most living reptiles the crocodiles, alligators, snakes, and lizards. 12

13 The euryapsids: have a single opening high on the skull, above the postorbital and squamosal bones, a condition derived from their diapsid ancestor. Include the ichthyosaurs and plesiosaurs. Most euryapsids had an aquatic or semiaquatic way of life. Thecodonts: important diapsid group which gave rise to the groups known as dinosaurs before the close of the Triassic. The thecodonts first appeared during the Early Triassic and became extinct at the close of the system. Some thecodonts ran on all four legs (quadrupedal), others exhibited a new bipedal, or two legged stance. By the end of the Triassic, thecodonts had given rise to both groups of dinosaurs, the Saurischia and Ornithischia. The Saurischia (lizard hipped): the termcomes from sauria, meaning reptile, and ischia, referring to the ischium bone of the pelvis. These dinosaurs had a pelvis built like that of many other reptiles, hence the name lizardhipped dinosaurs.from them evolved the other major group of dinosaurs. The Ornithischia (bird hipped):which appeared at the end of the Triassic. This group had a birdlike pelvis. TYPES OF REPTILES Different subclasses are distinguished by skull structure Anapsids: no holes on side of skull (turtles, extinct primitive [stem] reptiles) Synapsids: 1 hole on lower side of skull (extinct mammal like [pelycosaur & therapsid] reptiles). Synapsids ('fused arch'), also known as theropsids ('beast face'), are a class of animals that includes mammals and everything closer to mammals than to other living amniotes.the non mammalian members were traditionally described as mammallike reptiles, and are sometimes referred to as "proto mammals" or "stem mammals". Euryapsids: 1 hole on upper side of skull (most extinct marine reptiles. Examples of euryapsids are: icthyosaur, plesiosaur, nothosaur, placodont. This group of reptiles is included in the informal class Euryapsida which is considered to be an unnatural, polyphyletic group, as the various members are not closely related. This group of reptiles is distinguished by a single temporal fenestra, an opening behind the orbit, under which the post orbital and squamosal bones articulate. Diapsids: 2 HOLES on side of skull (MOST LIVING REPTILES [lizard, snake, crocodile, tuatara]; extinct DINOSAURS; extinct FLYING [pterosaur] REPTILES, & MOSASAURS [extinct marine reptiles]) Many new reptile types appeared in the Triassic TURTLES (Mesozoic turtles had teeth); TUATARAS: lizards of New Zealand CROCODILES (Cretaceous forms got to be as long as 15 m) LIZARDS & SNAKES (snakes evolved from lizards by the Cretaceous) MARINE & FLYING REPTILES & DINOSAURS (see below) THECODONTS (teeth set in sockets) a MOSTLY BIPEDAL GROUP that GAVE RISE to CROCODILES, LIZARDS, SNAKES, FLYING REPTILES & DINOSAURS The Triassic land fauna was dominated by therapsids & thecodonts MARINE REPTILES Several groups with marine adaptations, including paddle shaped limbs, streamlined bodies, & reproductive adaptations for birth of young at sea. Euryapsids:PLACODONTS; NOTHOSAURS; PLESIOSAURS (long necks; short, tailless bodies; flippers); ICTHYOSAURS (MOST FISH LIKE MARINE REPTILE [convergent with dolphins]) Anapsids:SEA TURTLES 13

14 Diapsids: MARINE CROCODILES OCCURRED MOSTLY DURING JURASSIC; MOSASAURS (SHORT NECKS; LONG BODIES & TAILS) Diapsid Skull: The name Diapsida means "two arches", and diapsids are traditionally classified based on their two ancestral skull openings (temporal fenestrae) posteriorly above and below the eye. Synapsid Skull: Synapsids evolved a temporal fenestra behind each eye orbit on the lateral surface of the skull. It may have evolved to provide new attachment sites for jaw muscles. Anapsid Skull: An anapsid is an amniote whose skull does not have openings near the temples Euryapsid Skull: distinguished by a single temporal fenestra, an opening behind the orbit, under which the post orbital and squamosal bones articulate. j: jugal p : parietal po : postorbital q : quadrate qj : quadratojugal sq : squamosal Images source: 14

15 Therapsids: "mammal like reptiles" cynodonts : ancestor of Mammalia. Thecodont ("socket toothed" reptile), now considered an obsolete term, was formerly used to describe a diverse range of early archosaurs that first appeared in the Latest Permian and flourished until the end of the Triassic period. The group includes the ancestors of dinosaurs (including birds), and ancestors of pterosaurs, and crocodilians, as well as a number of extinct forms that did not give rise to any descendants. Theropods Theropods are a group of bipedal saurischian dinosaurs. Although they were primarily carnivorous, a number of theropod families evolved herbivory during the Cretaceous System. Theropods first appear during the Carnian stage of the Upper Triassic about 220 million years ago (MYA) and were the sole large terrestrial carnivores from the Lower Jurassic until the close of the Cretaceous, about 65 MYA. Today, they are represented by the 9,300 living species of birds, which evolved in the Upper Jurassic from small specialized coelurosaurian dinosaurs. Tetrapoda: "four feet" thecodonts Archosauria (the "ruling reptiles": is a major group of diapsids, differentiated from the other diapsids by the presence of single openings in each side of the skull, in front of the eyes (antorbital fenestrae), among other characteristics. The ancestral archosaurs probably originated some 250 million years or so ago, in the upper Permian period. Their descendants (such as the dinosaurs) dominated the realm of the terrestrial vertebrates for a majority of the Mesozoic Era. Today, only the birds and crocodilians exist to provide a glimpse into the past glory of archosaurs. THE DINOSAURS 1. Evolved in the Triassic & expanded & diversified in the Jurassic & Cretaceous 2. Traditionally divided into two orders based on hip structure: Saurischia (lizard hipped) & Ornithischia (birdhipped) 3. Early dinosaurs were mostly Saurichians, which are divided into 2 suborders: (a) Bipedal carnivorous theropods, and (b) giant, 4 footed herbivorous sauropods 4. The Ornithischians are divided into 5 suborders:stegosaurs, Ankylosaurs, Ceratopsians, Ornithopods, and Pachycephalosauria All were herbivorous with the front teeth replaced by a beak & cheek teeth adapted for crushing coarse vegetation 15

16 Ornithopods, ankylosaours & ceratopsians were low browsers and were the dominant herbivores during the cretaceous Note: Robert Bakker has suggested that replacement of high browsing sauropods by low browsing ornithischians aided fast growing angiosperms in replacing slow growing gymnosperms as the dominant land plant Arguments in favor of warm bloodedness (Endothermy) Predator to prey ratios; erect stance; richly vascularized bones; growth rates; social behavior & migration of herds; hair on flying reptiles; complete dominance over mammals Arguments in favor of cold bloodedness (Ectothermy) Dinosaurs were reptiles & modern reptiles are cold blooded; erect stance & vascularized bones were responses to large size; large size itself Still an open question NOTE: it is no longer accepted that dinosaurs were slow & ponderous; dinosaur behavior probably like bird & mammal behavior. Theropods were quick & agile; large sauropods & ornithischians assembled in social herds; smaller sauropods & ornithischians behaved in a birdlike way. 16

17 SUMMARY OF DINOSAUR TAXONOMY Saurischia Ornithischia Therapoda Sauropoda Ornithopoda Pachycephalos auria Ankylosauria Stegosauria Allosaurus Compsognathus Deinonychus Tyrannosaurus Velociraptor Coelophysis Apatosaurus Brachiosaurus Camarasaurus Diplodocus Titanosaurus Hypsilophodon Iguanodon Parasaurolophus Stegoceras Ankylosaurus Stegosaurus al carnivores. Late Triassic to end of Cretaceous Size from 0.6 to 15 m Long, 2 or 3 kg to 7.3 metric tons. Some smaller genera may have hunted in packs. Quadrupedal herbivores. Late Triassic to Cretaceous, but most common during Jurassic. Size up to 27 m Long, 75 metric tons. Track ways indicate sauropods lived in herds. Preceded in fossil record by the smaller prosauropods ornithopods, such as Apatosaurus, had a billlike mouth and are called duck-billed dinosaurs. Size from a few meters Long up to 13 m and 3.6 metric tons. Especially diverse and common during the Cretaceous. Primarily bipedal herbivores, but could also walk on all fours. ceras only 2 m Long and 55 kg, but larger species known. Thick bones of skull cap might have aided in butting contests for dominance and mates. Bipedal herbivores of Cretaceous. osaurus more than 7 m Long and about 2.5 metric tons. Heavily armored with bony plates on top of head, back, and sides. Quadrupedal herbivore. ety of stegosaurs are known, but Stegosaurus, with bony 17

18 Ceratopsia Triceratops plates on its back and a spiked tail is best known. Plates probably were for absorbing and dissipating heat. Quadrupedal herbivores that were most common during the Jurassic. Stegosaurus 9 m Long, 1.8 metric tons. rous genera known. Some early ones bipedal, but Later Large animals were Quadrupedal herbivores. Much variation in size; Triceratops to 7.6 m Long and 5.4 metric tons, with Large bony frill over top of neck, three horns on skull, and beaklike mouth. Especially common during the Cretaceous. SAURISCHIAN DINOSAURS The saurischians include two distinct groups known as theropods and sauropods(see table above). Theropods(beast footed): Typical genus Tyrannosaurus;bipedal carnivores. Upper Triassic to end of Cretaceous.Size from 0.6 to 15 m long, 2 or 3 kg to 7.3 metric tons. Some smaller genera may have hunted in packs. All theropods were carnivorous bipeds that varied from tiny Compsognathus to giants such as Tyrannosaurus and similar but even larger species. Other genera include Allosaurus, Compsognathus, Deinonychus, Velociraptor, and Coelophysis. In 1996, Chinese scientists discovered several theropods with feathers. No one doubts that these dinosaurs had feathers, and molecular evidence indicates they were composed of the same material as bird feathers. Sauropods (reptile footed): Typical genus Brachiosaurus; Giant Quadrupedal herbivores. Upper Triassic to Cretaceous, but most common during Jurassic. Size up to 27 m long, 75 metric tons. Track ways indicate sauropods lived in herds. Preceded in fossil record by the smaller prosauropods includes the truly giant, quadrupedal herbivorous dinosaurs such as Apatosaurus, Diplodocus, Apatosaurus,Camarasaurus, and Titanosaurus; the largest land animals ever. Brachiosaurus, a giant even by sauropod standards, may have weighed 75 metric tons, and partial remains from several areas indicate that even larger sauropods may have existed. Track ways show that sauropods moved in herds. Sauropods were preceded in the fossil record by the smaller Upper Triassic to Lower Jurassic prosauropods, which were undoubtedly related to sauropods but probably not their ancestors. Sauropods were most common during the Jurassic; only a few genera existed during the Cretaceous. 18

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