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Lecture 13A: Paleozoic Life

Introduction

Proterozoic Setup

  • Single-celled organisms dominated Earth (Archean bacteria)

  • Early Proterozoic

    • Eukaryotic cells were a milestone in evolution.

  • Late Proterozoic

    • Exceptional preservation of soft-bodied organism that vaguely resemble jellyfish, sea pens, and wormlike animals.

    • Show older evolutionary history for our modern metazoans (multicellular animals).

Ediacaran Fauna

  • Neoproterozoic

  • Phylums Cnidaria (jellyfish, sea pens) and Echinodermata (starfish, sea urchins)

  • Fauna might go as far back as 800-900 Ma, but not enough evidence.

  • Widespread between 545-670 Ma.

Small Shelly Fauna

  • Very first mineralized elements in fossil record (600-550 Ma).

  • Preserved around the world (Esmeralda County, NV).

  • Fossils suggest biological pathways for secreting a mineral were in place before Cambrian.

  • Although mechanism to form shell was there, not really being used

Cloudina

  • Cloudina: One of earliest known animal fossil with mineralized skeleton.

  • Metazoan of Neoproterozoic (Ediacaran) to Cambrian.

  • Can be describe as a “tube-dwelling worm”.

  • First “shelly” critter.

Bore Holes in Shells

  • Implying there was at least some form of carnivore in Ediacaran.

    • Comes from predation.

  • Likely some sore of sponge

  • Sponges were the apex predator at the time (top of the food chain).

Non-Vertebrates

  • Non-vertebrate: Animals that do not have a backbone or spine. This group includes creatures like insects, worms, jellyfish, and snails.

  • Phylum: A major category in the classification of living organisms, grouping together related classes.

  • Porifera - the sponges

  • Cnidaria - corals, jellyfish, sea anemones

  • Arthropoda - insects, spider, trilobites

  • Mollusca - clams, oysters, snails, octopods

  • Echinodermata - sea urchins, star fish

Cambrian

Important Cambrian Biology

  • Cambrian Explosion (seemingly instantaneous appearance of many varied animals)

  • Trilobites (index fossil)

  • Treptichnus pedum (trace fossil)

  • Burgess Shale Lagerstatten (one of the major lagerstatten)

  • Anomalocaris (apex predator)

Defining the Cambrian

  • First period of Paleozoic Era and Phanerozoic Eon

  • Phanerozoic → ancient Greek, “visible life”

  • We now have a “visible” Ediacaran biota

  • Why is Ediacaran not a period of Phanerozoic?

    • Traditionally base of Cambrian was not defined by “visible life”.

    • It was defined by first occurrence of Trilobites.

Trilobites

  • Phylum: Arthropoda

  • Older and older trilobites kept being found, so the boundary kept having to be moved.

  • Good index fossil

Led to a Debate

  • Russian/Chinese scientist wanted Cloudina to define Cambrian.

  • U.S. scientists anted to keep Trilobite.

  • Canadians proposed a trace fossil.

    • Treptichnus pedum → preserved worm burrow.

    • Occurs above Cloudina but below Trilobites.

  • The treptichnus pedum defines the base of the Cambrian.

Cambrian Explosion

  • Cambrian Explosion: Refers to the seemingly instantaneous appearance of many varied animals in the early to middle Cambrian.

    • It’s more like a slow fuse, this life didn’t suddenly appear.

  • New phyla appears: Arthropoda, Chordata, Mollusca.

What’s Going On?

  • Changes attributed to Cambrian Explosion.

  • Hard parts are evolving as a defense against predation.

    • Justification: phenomenon occurs across several unrelated phyla same time thus, it must be a reaction to an ecological stress.

  • Sea water chemistry changed to the point that secreting a shell out of carbonate or phosphate was not metabolically overly expensive.

    • Justification: phenomenon occurs across several unrelated phyla same time thus, chemistry must have changed or else all these groups would have evolved hard parts sooner.

  • Take away → Cambrian explosion doesn’t really have a good explanation for why it happened when it did.

Lagerstatten

  • Lagerstatten: An environment where fossils are exceptionally well-preserved, often including soft tissues.

  • An instance of exceptional preservation.

  • Environments give us immaculate pictures of prehistoric life.

  • Some are preserved in three dimensions.

  • Many remains of animals lacked hard parts.

Burgess Shale

  • Discovered in 1909 on an expedition to collect Cambrian trilobites in the Canadian Rockies by Charles Walcott.

  • One of the major lagerstatten.

Lack of Oxygen (Burgess Preservation)

  • Mudslides and anoxic water (no oxygen)

Cambrian Life

  • Simple in structure, ecologically diverse.

  • Archaeocyathids and other reef building Porifera (sponges).

    • Archaeocyathids: An extinct group of ancient marine sponges that lived during the Cambrian period; some of the earliest known reef-building organisms.

  • Trilobites

  • Brachiopods, Mollusca, and others

Top Cambrian Predator

  • Top predator was the arthropod Anomalocaris.

  • “Odd shrimp”

  • Mouth plates used for crushing prey captured by these appendages.

  • Trilobites with bite marks in fossil record.

Evolutionary Patterns

  • Cambrian marks a time when Metazoans were ecologically experimenting.

  • The Cambrian marks diversification of Phyla, but Family / Genus/ Species diversity remained low.

  • Only 1 Phylum (Bryozoans) originated after Cambrian but Family diversity explodes in the subsequent Ordovician → GOBE

Ordovician

Important Ordovician Biology

  • Great Ordovician Biodiversification Event (GOBE) (tripling of global diversity)

  • Coral reefs (Rugose and Tabulates)

  • Nautiloids and sea stars (top predators of the Ordivician)

  • Vertebrate radiation (earliest = jawless fish; reptiles, mammal, fish, birds, and amphibians)

  • Ordovician mass extinction event (first great Mass Extinction; caused by global cooling)

GOBE

  • Great Ordovician Biodiversification Event

  • Mid-Ordovician (over ~25 million years geologic time)

  • Unlike Cambrian diversification (new phyla), this was characterized by an increase in species, genus, family, and order.

  • Tripling of global diversity.

  • Consistent throughout Paleozoic.

  • Simple Cambrian food chain of deposit and suspension feeders replaced by more complex food web.

Causes of Ordovician Radiation

  • Late Cambrian extinctions left niches (function) available.

  • A time of continental fragmentation and sea-floor spreading.

    • Which may have provided nutrients

  • High sea levels created vast shallow warm seas (Tippecanoe).

  • Oxygen levels reaching modern levels.

  • Evolution of new predators

    • Robust calcite shells evolve

Brachiopods

  • During Paleozoic, thousands of different species in near-shore and deep-sea.

  • Found in Cambrian but became abundant in Ordovician (calcite becoming more commonplace).

  • Useful for biostratigraphic correlation.

Bryozoans

  • First show up in the Ordovician.

  • Filter feeding “Moss Animals”

    • Colonial organism which supported 100’s of little animals.

    • Build coral-like structures.

    • Cousins of the brachiopods

Crinoids

  • Known as “Sea lilies”

  • Stalked echinoderms (phylum includes sea stars, sea urchins, etc.)

  • Sessile (immobile); modern ones are mobile.

  • Arms used as long filter-feeding fans.

  • Bodies suspended by long stalks rooted to sea bed.

    • Rarely preserved

Corals

  • Sponge reefs occurred during Cambrian.

  • Ordovician marks first coral reefs.

    • Rugose Corals: Horn corals

    • Tabulates: Honeycomb corals

Stromatoporoids

  • Extinct group of mound-building (reef) sponges.

  • Common during Ordovician-Devonian

  • Difficult to interpret (enigmatic); used to think these were stromatolites, but these are animals, not cyanobacteria.

  • Grew in association with corals, brachiopods, and other reef-dwellers.

Top Predator(s)

  • Cambrian: largest predator arthropod Anomalocaris.

  • Ordovician: Cephalopod nautiloids reached up to 10 m.

    • Living relative is the Chambered nautilus

  • The other new predator were sea stars.

Gastropods

  • Evolve in Cambrian

  • Snails

  • Predators of the stromatolites

  • Diversify in the Ordovician

    • Show abilities to scavenge

    • Predation

    • Graze (results in decline of stromatolites)

Graptolites

  • Ordovician index fossils

  • Greek for “writing in the rocks”

  • Colonial animals composed of interconnected system of tubes.

  • Filter feeders

  • Extinct

Vertebrates

  • Ordovician marks the biodiversification (or radiation) of our own subphylum → the Vertebrates.

    • Fossil discoveries (in China) can place back to the Cambrian.

  • Earliest vertebrates were jawless fish.

  • Reptiles, mammals, fishes, birds, amphibians

Agnathans

  • Ancient Greek for “no jaws”

  • Oldest known discovery was in the Lower Cambrian, but diversification in Ordovician.

  • Relatives exist today.

Conodonts

  • Class of agnathans

  • Group of well-known, small, tooth-like fossils composed of the mineral apatite (calcium-phosphate).

  • Evolve in Cambrian but become abundant in the Ordovician

  • Become excellent index fossils for the Ordovician.

  • Extinct at end of Triassic

First Life on Land

  • Seeing small signs that life existed on land.

  • Life invades land

    • Land plants ~475 Ma

    • First land colonizers were likely micro-biotic crusts.

      • Communities of fungi, bacteria, and algae.

  • Advantages:

    • Light abundant (photosynthesis)

    • Nutrients supplied directly to soils (weathering).

  • First fossils of spores and tissues of simple land plants in later Ordovician.

Ordovician Extinction

  • The first great Mass Extinction

  • Global cooling caused the Ordovician mass extinction event.

  • >100 families of marine animals did not make it to Silurian.

  • Half of all brachiopods and bryozoans died out.

  • Reef communities decimated; did not bounce back until late Silurian.

  • Nautiloids were decimated.

  • Trilobites declined further

  • Likely cause was a global glaciation event

    • Created a dramatic drop in sea-level eliminating much habitat along continental shelved.

  • Concentrated on tropical groups (most warm water invertebrates were gone).

  • Cold or deep water adapted groups weathered the storm best.

  • Early Silurian Seas were dominated by low diversity widespread taxa; by the Late Silurian and into the Devonian, life had recovered.


Review Questions (All of Lecture 13)

  • Were there general names for specific periods and/or eras (e.g., Age of Fish)?

    • Age of Fish: Devonian

  • How did life change from Precambrian into (and through) the Paleozoic?

    • Life evolved from simple, single-celled organisms in the Precambrian to more complex, multicellular forms in the Paleozoic.

    • The Cambrian Explosion led to a rapid increase in biodiversity and the evolution of major animal groups.

    • By the end of the Paleozoic, life had colonized land, but the era ended with a mass extinction.

  • What was the Cambrian explosion?

    • Cambrian Explosion: Refers to the seemingly instantaneous appearance of many varied animals in the early to middle Cambrian.

    • New phyla appears: Arthropoda, Chordata, Mollusca.

  • What is a Lagerstatten? Why was the Burgess Shale important? Who discovered it?

    • Lagerstatten: An environment where fossils are exceptionally well-preserved, often including soft tissues.

    • Burgess Shale is one of the major lagerstatten.

    • Discovered by Charles Walcott on an expedition to collect Cambrian trilobites in the Canadian Rockies.

  • What were dominant predators, new organisms, and index fossils for specific Periods?

    • Paleozoic:

      • Apex predator: Sponge

    • Cambrian:

      • Apex predator: Anomalocaris

      • Index fossil: Trilobites

      • Trace fossil: Treptichnus pedum

    • Ordovician:

      • Apex predator: Cephalopod

      • Index fossil: Graptolites, conodonts

  • What was the evolutionary path of life from marine to land?

    • Early signs of life on land.

    • Life invades land with plants around 475 Ma.

    • Microbiotic crusts (fungi, bacteria, algae) were likely the first colonizers.

    • Advantages: abundant light for photosynthesis and nutrients from soil weathering.

    • First fossils of simple land plants appear in the late Ordovician.

  • What were the mass extinction event(s)? When did they occur? What caused them?

    • Ordovician mass extinction event

      • First great mass extinction

      • Global glaciation event (global cooling)

S

Lecture 13A: Paleozoic Life

Introduction

Proterozoic Setup

  • Single-celled organisms dominated Earth (Archean bacteria)

  • Early Proterozoic

    • Eukaryotic cells were a milestone in evolution.

  • Late Proterozoic

    • Exceptional preservation of soft-bodied organism that vaguely resemble jellyfish, sea pens, and wormlike animals.

    • Show older evolutionary history for our modern metazoans (multicellular animals).

Ediacaran Fauna

  • Neoproterozoic

  • Phylums Cnidaria (jellyfish, sea pens) and Echinodermata (starfish, sea urchins)

  • Fauna might go as far back as 800-900 Ma, but not enough evidence.

  • Widespread between 545-670 Ma.

Small Shelly Fauna

  • Very first mineralized elements in fossil record (600-550 Ma).

  • Preserved around the world (Esmeralda County, NV).

  • Fossils suggest biological pathways for secreting a mineral were in place before Cambrian.

  • Although mechanism to form shell was there, not really being used

Cloudina

  • Cloudina: One of earliest known animal fossil with mineralized skeleton.

  • Metazoan of Neoproterozoic (Ediacaran) to Cambrian.

  • Can be describe as a “tube-dwelling worm”.

  • First “shelly” critter.

Bore Holes in Shells

  • Implying there was at least some form of carnivore in Ediacaran.

    • Comes from predation.

  • Likely some sore of sponge

  • Sponges were the apex predator at the time (top of the food chain).

Non-Vertebrates

  • Non-vertebrate: Animals that do not have a backbone or spine. This group includes creatures like insects, worms, jellyfish, and snails.

  • Phylum: A major category in the classification of living organisms, grouping together related classes.

  • Porifera - the sponges

  • Cnidaria - corals, jellyfish, sea anemones

  • Arthropoda - insects, spider, trilobites

  • Mollusca - clams, oysters, snails, octopods

  • Echinodermata - sea urchins, star fish

Cambrian

Important Cambrian Biology

  • Cambrian Explosion (seemingly instantaneous appearance of many varied animals)

  • Trilobites (index fossil)

  • Treptichnus pedum (trace fossil)

  • Burgess Shale Lagerstatten (one of the major lagerstatten)

  • Anomalocaris (apex predator)

Defining the Cambrian

  • First period of Paleozoic Era and Phanerozoic Eon

  • Phanerozoic → ancient Greek, “visible life”

  • We now have a “visible” Ediacaran biota

  • Why is Ediacaran not a period of Phanerozoic?

    • Traditionally base of Cambrian was not defined by “visible life”.

    • It was defined by first occurrence of Trilobites.

Trilobites

  • Phylum: Arthropoda

  • Older and older trilobites kept being found, so the boundary kept having to be moved.

  • Good index fossil

Led to a Debate

  • Russian/Chinese scientist wanted Cloudina to define Cambrian.

  • U.S. scientists anted to keep Trilobite.

  • Canadians proposed a trace fossil.

    • Treptichnus pedum → preserved worm burrow.

    • Occurs above Cloudina but below Trilobites.

  • The treptichnus pedum defines the base of the Cambrian.

Cambrian Explosion

  • Cambrian Explosion: Refers to the seemingly instantaneous appearance of many varied animals in the early to middle Cambrian.

    • It’s more like a slow fuse, this life didn’t suddenly appear.

  • New phyla appears: Arthropoda, Chordata, Mollusca.

What’s Going On?

  • Changes attributed to Cambrian Explosion.

  • Hard parts are evolving as a defense against predation.

    • Justification: phenomenon occurs across several unrelated phyla same time thus, it must be a reaction to an ecological stress.

  • Sea water chemistry changed to the point that secreting a shell out of carbonate or phosphate was not metabolically overly expensive.

    • Justification: phenomenon occurs across several unrelated phyla same time thus, chemistry must have changed or else all these groups would have evolved hard parts sooner.

  • Take away → Cambrian explosion doesn’t really have a good explanation for why it happened when it did.

Lagerstatten

  • Lagerstatten: An environment where fossils are exceptionally well-preserved, often including soft tissues.

  • An instance of exceptional preservation.

  • Environments give us immaculate pictures of prehistoric life.

  • Some are preserved in three dimensions.

  • Many remains of animals lacked hard parts.

Burgess Shale

  • Discovered in 1909 on an expedition to collect Cambrian trilobites in the Canadian Rockies by Charles Walcott.

  • One of the major lagerstatten.

Lack of Oxygen (Burgess Preservation)

  • Mudslides and anoxic water (no oxygen)

Cambrian Life

  • Simple in structure, ecologically diverse.

  • Archaeocyathids and other reef building Porifera (sponges).

    • Archaeocyathids: An extinct group of ancient marine sponges that lived during the Cambrian period; some of the earliest known reef-building organisms.

  • Trilobites

  • Brachiopods, Mollusca, and others

Top Cambrian Predator

  • Top predator was the arthropod Anomalocaris.

  • “Odd shrimp”

  • Mouth plates used for crushing prey captured by these appendages.

  • Trilobites with bite marks in fossil record.

Evolutionary Patterns

  • Cambrian marks a time when Metazoans were ecologically experimenting.

  • The Cambrian marks diversification of Phyla, but Family / Genus/ Species diversity remained low.

  • Only 1 Phylum (Bryozoans) originated after Cambrian but Family diversity explodes in the subsequent Ordovician → GOBE

Ordovician

Important Ordovician Biology

  • Great Ordovician Biodiversification Event (GOBE) (tripling of global diversity)

  • Coral reefs (Rugose and Tabulates)

  • Nautiloids and sea stars (top predators of the Ordivician)

  • Vertebrate radiation (earliest = jawless fish; reptiles, mammal, fish, birds, and amphibians)

  • Ordovician mass extinction event (first great Mass Extinction; caused by global cooling)

GOBE

  • Great Ordovician Biodiversification Event

  • Mid-Ordovician (over ~25 million years geologic time)

  • Unlike Cambrian diversification (new phyla), this was characterized by an increase in species, genus, family, and order.

  • Tripling of global diversity.

  • Consistent throughout Paleozoic.

  • Simple Cambrian food chain of deposit and suspension feeders replaced by more complex food web.

Causes of Ordovician Radiation

  • Late Cambrian extinctions left niches (function) available.

  • A time of continental fragmentation and sea-floor spreading.

    • Which may have provided nutrients

  • High sea levels created vast shallow warm seas (Tippecanoe).

  • Oxygen levels reaching modern levels.

  • Evolution of new predators

    • Robust calcite shells evolve

Brachiopods

  • During Paleozoic, thousands of different species in near-shore and deep-sea.

  • Found in Cambrian but became abundant in Ordovician (calcite becoming more commonplace).

  • Useful for biostratigraphic correlation.

Bryozoans

  • First show up in the Ordovician.

  • Filter feeding “Moss Animals”

    • Colonial organism which supported 100’s of little animals.

    • Build coral-like structures.

    • Cousins of the brachiopods

Crinoids

  • Known as “Sea lilies”

  • Stalked echinoderms (phylum includes sea stars, sea urchins, etc.)

  • Sessile (immobile); modern ones are mobile.

  • Arms used as long filter-feeding fans.

  • Bodies suspended by long stalks rooted to sea bed.

    • Rarely preserved

Corals

  • Sponge reefs occurred during Cambrian.

  • Ordovician marks first coral reefs.

    • Rugose Corals: Horn corals

    • Tabulates: Honeycomb corals

Stromatoporoids

  • Extinct group of mound-building (reef) sponges.

  • Common during Ordovician-Devonian

  • Difficult to interpret (enigmatic); used to think these were stromatolites, but these are animals, not cyanobacteria.

  • Grew in association with corals, brachiopods, and other reef-dwellers.

Top Predator(s)

  • Cambrian: largest predator arthropod Anomalocaris.

  • Ordovician: Cephalopod nautiloids reached up to 10 m.

    • Living relative is the Chambered nautilus

  • The other new predator were sea stars.

Gastropods

  • Evolve in Cambrian

  • Snails

  • Predators of the stromatolites

  • Diversify in the Ordovician

    • Show abilities to scavenge

    • Predation

    • Graze (results in decline of stromatolites)

Graptolites

  • Ordovician index fossils

  • Greek for “writing in the rocks”

  • Colonial animals composed of interconnected system of tubes.

  • Filter feeders

  • Extinct

Vertebrates

  • Ordovician marks the biodiversification (or radiation) of our own subphylum → the Vertebrates.

    • Fossil discoveries (in China) can place back to the Cambrian.

  • Earliest vertebrates were jawless fish.

  • Reptiles, mammals, fishes, birds, amphibians

Agnathans

  • Ancient Greek for “no jaws”

  • Oldest known discovery was in the Lower Cambrian, but diversification in Ordovician.

  • Relatives exist today.

Conodonts

  • Class of agnathans

  • Group of well-known, small, tooth-like fossils composed of the mineral apatite (calcium-phosphate).

  • Evolve in Cambrian but become abundant in the Ordovician

  • Become excellent index fossils for the Ordovician.

  • Extinct at end of Triassic

First Life on Land

  • Seeing small signs that life existed on land.

  • Life invades land

    • Land plants ~475 Ma

    • First land colonizers were likely micro-biotic crusts.

      • Communities of fungi, bacteria, and algae.

  • Advantages:

    • Light abundant (photosynthesis)

    • Nutrients supplied directly to soils (weathering).

  • First fossils of spores and tissues of simple land plants in later Ordovician.

Ordovician Extinction

  • The first great Mass Extinction

  • Global cooling caused the Ordovician mass extinction event.

  • >100 families of marine animals did not make it to Silurian.

  • Half of all brachiopods and bryozoans died out.

  • Reef communities decimated; did not bounce back until late Silurian.

  • Nautiloids were decimated.

  • Trilobites declined further

  • Likely cause was a global glaciation event

    • Created a dramatic drop in sea-level eliminating much habitat along continental shelved.

  • Concentrated on tropical groups (most warm water invertebrates were gone).

  • Cold or deep water adapted groups weathered the storm best.

  • Early Silurian Seas were dominated by low diversity widespread taxa; by the Late Silurian and into the Devonian, life had recovered.


Review Questions (All of Lecture 13)

  • Were there general names for specific periods and/or eras (e.g., Age of Fish)?

    • Age of Fish: Devonian

  • How did life change from Precambrian into (and through) the Paleozoic?

    • Life evolved from simple, single-celled organisms in the Precambrian to more complex, multicellular forms in the Paleozoic.

    • The Cambrian Explosion led to a rapid increase in biodiversity and the evolution of major animal groups.

    • By the end of the Paleozoic, life had colonized land, but the era ended with a mass extinction.

  • What was the Cambrian explosion?

    • Cambrian Explosion: Refers to the seemingly instantaneous appearance of many varied animals in the early to middle Cambrian.

    • New phyla appears: Arthropoda, Chordata, Mollusca.

  • What is a Lagerstatten? Why was the Burgess Shale important? Who discovered it?

    • Lagerstatten: An environment where fossils are exceptionally well-preserved, often including soft tissues.

    • Burgess Shale is one of the major lagerstatten.

    • Discovered by Charles Walcott on an expedition to collect Cambrian trilobites in the Canadian Rockies.

  • What were dominant predators, new organisms, and index fossils for specific Periods?

    • Paleozoic:

      • Apex predator: Sponge

    • Cambrian:

      • Apex predator: Anomalocaris

      • Index fossil: Trilobites

      • Trace fossil: Treptichnus pedum

    • Ordovician:

      • Apex predator: Cephalopod

      • Index fossil: Graptolites, conodonts

  • What was the evolutionary path of life from marine to land?

    • Early signs of life on land.

    • Life invades land with plants around 475 Ma.

    • Microbiotic crusts (fungi, bacteria, algae) were likely the first colonizers.

    • Advantages: abundant light for photosynthesis and nutrients from soil weathering.

    • First fossils of simple land plants appear in the late Ordovician.

  • What were the mass extinction event(s)? When did they occur? What caused them?

    • Ordovician mass extinction event

      • First great mass extinction

      • Global glaciation event (global cooling)