The Cambrian Explosion: Unveiling the Early History of Life through the Burgess Shale

1. Chapter 12 Paleozoic Life History: Invertebratesmain points this chapter-1. animals with skeletons appear in Cambrian2. ecosystem interacts with environment3. Cambrian-evolutionary innovations4. Ordovician- dramatic increase in diversity, mass extinction at end Ordovician5. Silurian& Devonian: re diversificationafter end Ord mass extinction; major reef building6. Late Devonian extinctions: marine community again re diversified, with new organisms7. end of Permian- greatest mass extinction in Earth’s history occurred

2. Study of Paleozoic Life • We will examine the history of Paleozoic life • as a system of interconnected biologic and geologic events • Evolution and plate tectonics • are the forces that drove this system • The opening and closing of ocean basins, • transgressions and regressions of epeiric seas, • the formation of mountain ranges, • and the changing positions of the continents • had a profound effect on the evolution • of the marine and terrestrial communities

3. The Cambrian Explosion • At the beginning of the Paleozoic Era, • animals with skeletons • appeared rather abruptly in the fossil record • In fact, their appearance is described • as an explosive development • of new types of animals • and is referred to as • the "Cambrian explosion" by most scientists • It is rapid, however, only in the context of geologic time, having taken place over millions of years

4. Burgess Shale Soft-Bodied Fossils • On August 30 and 31, 1909, • Charles D. Walcott, • geologist and head of the Smithsonian Institution, • discovered the first soft-bodied fossils • from the Burgess Shale, • a discovery of immense importance in deciphering the early history of life • Walcott and his collecting party split open numerous blocks of shale, • many of which yielded the impressions • of a number of soft-bodied organisms • beautifully preserved on bedding planes

5. More Complete Picture of a Middle Cambrian Community • The importance of Walcott's discovery • is that it allowed geologists a rare glimpse into a world previously almost unknown • that of the soft-bodied animals that lived some 530 million years ago • The beautifully preserved fossils • from the Burgess Shale • present a much more complete picture • of a Middle Cambrian community • than deposits containing only fossils of the hard parts of organisms

6. Burgess Shale • Diorama of the environment and biota • of the Phyllopod bed of the Burgess Shale, • British Columbia, Canada • algae • sponges • among others

7. Sixty Percent Soft-Bodied • In fact, 60% of the total fossil assemblage • of more than 100 genera is composed of soft-bodied animals, • a percentage comparable to present-day marine communities • What conditions led to the remarkable preservation of the Burgess Shale fauna? • The site of deposition of the Burgess Shale • was located at the base of a steep submarine escarpment

8. Reason for the Preservation • The animals • whose exquisitely preserved fossil remains • are found in the Burgess Shale • lived in and on mud banks • that formed along the top of this escarpment • Periodically, this unstable area • would slump and slide down the escarpment • as a turbidity current • At the base, the mud and animals carried with it • were deposited in a deep-water anaerobic environment devoid of life

9. Carbonaceous Impressions • In such an environment, • bacterial degradation did not destroy the buried animals • and they were compressed by the weight of the overlying sediments • and eventually preserved as carbonaceous impressions

10. Rare Preservation: Burgess Shale • Ottoia, a carnivorous worm

11. Rare Preservation: Burgess Shale • Wiwaxia, a scaly armored sluglike creature whose affinities remain controversial

12. Rare Preservation: Burgess Shale • Hallucigenia, a velvet worm

13. Rare Preservation: Burgess Shale • Waptia, an anthropod

14. Basic Body Plans • These were followed by • an explosion of invertebrate phyla • during the Cambrian, • some of which are now extinct • These Cambrian phyla • represent the rootstock • and basic body plans • from which all present-day invertebrates evolved

15. Strangeness of the Burgess Shale Biota • In other words, life was much more diverse • in terms of phyla • during the Cambrian • than it is today • The reason members of the Burgess Shale biota • look so strange to us • is that no living organisms • possess their basic body plan, • and therefore many of them have been placed into new phyla

16. Triggering Mechanism • It appears likely that the Cambrian explosion • probably had its roots firmly planted in the Proterozoic • However, the mechanism • that triggered this event is still unknown and • was likely a combination of factors, • both biological and geological • For example, geologic evidence • indicates Earth was glaciated • one or more times during the Proterozoic, • followed by global warming during the Cambrian

17. Major Event in Earth's History • Whatever the ultimate cause of the Cambrian explosion, • the appearance of a skeletonized fauna • and the rapid diversification of that fauna • during the Early Cambrian • was a major event in Earth's history

18. Sharp Contrast • The sudden appearance of shelled animals • during the Early Cambrian • contrasts sharply with the biota living • during the preceding Proterozoic Eon • Up until the evolution of the Ediacaran fauna, • Earth was populated primarily • by single-celled organisms • The Ediacaran fauna, • which is found on all continents except Antarctica, • consists primarily of multicelled soft-bodied organisms

19. Lower Cambrian Shelly Fossil • The tube of an anabaritid from the Mackenzie Mountains, Northwest Territories, Canada • This specimen is several millimeters in size • Archaeooides, an enigmatic spherical • fossil from the Mackenzie Mountains, • Northwest Territories, Canada • A conical sclerite* of • Lapworthella from Australia • a piece of the armor covering

20. Why Skeletons • Along with the question of • why did animals appear so suddenly in the fossil record • is the equally intriguing one of • why they initially acquired skeletons • and what selective advantage this provided • A variety of explanations • about why marine organisms evolved skeletons • have been proposed, • but none is completely satisfactory or universally accepted

21. Advantages of an Exoskeleton • The formation of an exoskeleton • confers many advantages on an organism: (1) It provides protection against ultraviolet radiation, allowing animals to move into shallower waters; (2) it helps prevent drying out in an intertidal environment; (3) it provides protection against predators • Recent evidence of actual fossils of predators • and specimens of damaged prey, • as well as antipredatory adaptations in some animals, • indicates that the impact of predation during the Cambrian was great

22. Cambrian Predator • Reconstruction of Anamalocaris • a predator from the Early and Middle Cambrian • It was about 45 cm long and probably fed on trilobites • Its gripping appendages presumably carried food to its mouth

23. Wounded Trilobite • Wounds to the body of the trilobite Olenellus robsonensis • The wounds have healed, demonstrating that they occurred when the animal was alive and were not inflicted on an empty shell

24. Advantages of an Exoskeleton • With predators playing an important role • in the Cambrian marine ecosystem, • any mechanism or feature • that protected an animal • would certainly be advantageous • and confer an adaptive advantage to the organism (4) A fourth advantage is that • a supporting skeleton, whether an exo- or endoskeleton, • allows animals to increase their size • and provides attachment sites for muscles

25. It Is Unknown Why Organisms Evolved Mineralized Skeletons • There currently is no clear answer about • why marine organisms evolved mineralized skeletons • during the Cambrian explosion and shortly thereafter • They undoubtedly evolved • because of a variety of biologic and environmental factors

26. Mineralized Skeletons Were Successful • Whatever the reason, • the acquisition of a mineralized skeleton • was a major evolutionary innovation • allowing invertebrates to successfully occupy • a wide variety of marine habitats

27. Marine Invertebrate Communities • Rather than focusing on • the history of each invertebrate phylum, • we will survey the evolution • of the marine invertebrate communities through time, • concentrating on the major features and changes that took place • To do that, we need to briefly examine • the nature and structure • of living marine communities so that • we can make a reasonable interpretation • of the fossil record

28. The Present Marine Ecosystem • In analyzing the present-day marine ecosystem, • we must look at where organisms live, • how they get around, • as well as how they feed • Organisms that live in the water column • above the seafloor • are called pelagic • They can be divided into two main groups: • the floaters, or plankton, • and the swimmers, or nekton

29. Plankton • Plankton are mostly passive and go where currents carry them • Plant plankton • such as diatoms, dinoflagellates, and various algae, • are called phytoplankton and are mostly microscopic • Animal plankton are called zooplankton and are also mostly microscopic • Examples of zooplankton include foraminifera, radiolarians, and jellyfish

30. Nekton • The nekton are swimmers • and are mainly vertebrates • such as fish; • the invertebrate nekton • include cephalopods

31. Benthos • Organisms that live • on or in the seafloor make up the benthos • They can be characterized • as epifauna (animals) or epiflora (plants), • for those that live on the seafloor, • or as infauna, • which are animals living in and moving through the sediments

32. Sessile and Mobile • The benthos can be further divided • into those organisms that stay in one place, • called sessile, • and those that move around on or in the seafloor, • called mobile

33. Marine Ecosystem-Where and how animals and plants live in the marine ecosystem Plankton: Jelly fish Sessile epiflora: seaweed Nekton: fish cephalopod Sessile epifauna: bivalve Benthos: d-k crinoid coral

34. Marine Ecosystem Infauna: worm, bivalve Mobile epifauna: gastropod, starfish

35. Feeding Strategies • The feeding strategies of organisms • are also important in terms of their relationships • with other organisms in the marine ecosystem • There are basically four feeding groups: • suspension-feeding animals remove or consume microscopic plants and animals as well as dissolved nutrients from the water; • herbivores are plant eaters; • carnivore-scavengers are meat eaters; • and sediment-deposit feeders ingest sediment and extract the nutrients from it

36. Marine Ecosystem coral crinoid bivalve Suspension feeders:

37. Marine Ecosystem worm sediment-deposit feeder Herbivores: gastropod Carnivores-scavengers: starfish

38. Organism's Place • We can define an organism's place • in the marine ecosystem • by where it lives • and how it eats • For example, an articulate brachiopod • is a benthonic, • epifaunal suspension feeder, • whereas a cephalopod • is a nektonic carnivore

39. Trophic Levels • An ecosystem includes several trophic levels, • which are tiers of food production and consumption • within a feeding hierarchy • The feeding hierarchy • and hence energy flow • in an ecosystem comprise • a food web of complex interrelationships among • the producers, • consumers, • and decomposers

40. Primary Producers • The primary producers, or autotrophs, • are those organisms that manufacture their own food • Virtually all marine primary producers are phytoplankton • Feeding on the primary producers • are the primary consumers, which are mostly suspension feeders

41. Other Consumers • Secondary consumers feed on • the primary consumers, • and thus are predators, while tertiary consumers, which are also predators, feed on the secondary consumers • Besides the producers and consumers, • there are also transformers and decomposers • These are bacteria that break down the dead organisms • that have not been consumed • into organic compounds that are then recycled

42. Marine Food Web • Showing the relationships • among the • producers, • consumers, • and decomposers

43. When the System Changes • When we look at the marine realm today, • we see a complex organization of organisms • interrelated by trophic interactions • and affected by changes in the physical environment • When one part of the system changes, • the whole structure changes, • sometimes almost insignificantly, • other times catastrophically

44. Changing Marine Ecosystem • As we examine the evolution of the Paleozoic marine ecosystem, • keep in mind how geologic and evolutionary changes • can have a significant impact on its composition and structure • For example, the major transgressions onto the craton • opened up vast areas of shallow seas • that could be inhabited • The movement of continents • affected oceanic circulation patterns • as well as causing environmental changes This one slide says it all…..

45. Cambrian Marine Community • The Cambrian Period was a time • during which many new body plans evolved • and animals moved into new niches • As might be expected, the Cambrian • witnessed a higher percentage of such experiments • than any other period of geologic history

46. Cambrian Skeletonized Life • Although almost all the major invertebrate phyla • evolved during the Cambrian Period • many were represented by only a few species • While trace fossils are common • and echinoderms diverse, • the organisms that comprised the majority of Cambrian skeletonized life were • trilobites, • inarticulate brachiopods, • and archaeocyathids

47. Cambrian Marine Community • Floating jellyfish, swimming arthropods, benthonic sponges, and scavenging trilobites Reconstruction

48. Trilobites • Trilobites were • by far the most conspicuous element • of the Cambrian marine invertebrate community • and made up about half of the total fauna • Trilobites were • benthonic • mobile • sediment-deposit feeders • that crawled or swam along the seafloor

49. Trilobites • They first appeared in the Early Cambrian, • rapidly diversified, • reached their maximum diversity • in the Late Cambrian, • and then suffered mass extinctions • near the end of the Cambrian • from which they never fully recovered • As yet no consensus exists on what caused the trilobite extinctions,

50. Trilobite Extinctions • but a combination of factors were likely involved, • including possibly a reduction of shelf space, • increased competition, • and a rise in predators • It has also been suggested • that a cooling of the seas may have played a role, • particularly for the extinctions • that took place • at the end of the Ordovician Period