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NJU Course Extinctions: Past, Present & Future

Prof. Norman MacLeod School of Earth Sciences & Engineering, Nanjing University Extinctions: Past, Present & Future Extinctions: Past, Present & Future

Course Syllabus (Revised)

Section Week Title Introduction 1 Course Introduction, Intro. To Extinction Introduction 2 History of Extinction Studies Introduction 3 Evolution, Fossils, Time & Extinction Extinctions 4 Origin of Life & Precambrian Extionctions Extinctions 5 Early Paleozoic World & Extinctions Paleozoic Extinctions 6 Middle Paleozoic World & Extinctions Paleozoic Extinctions 7 Late Paleozoic World & Extinctions Assessment 8 Mid-Term Examination Extinctions 9 - World & Extinctions Mesozoic Extinctions 10 Labor Day Holiday Cenozoic Extinctions 11 World & Extinctions Cenozoic Extinctions 12 Paleogene World & Extinctions Cenozoic Extinctions 13 Neogene World & Extinctions Modern Extinctions 14 Quaternary World & Extinctions Modern Extinctions 15 Modern World: Floras, Faunas & Environment Modern Extinctions 16 Modern World: Habitats & Organisms Assessment 17 Final Examination Early Paleozoic World, Life & Extinctions Norman MacLeod School of Earth Sciences & Engineering, Nanjing University Early Paleozoic World, Life & Extinctions

Objectives

Understand the structure of the early Paleozoic world in terms of timescales, geography, environ- ments, and organisms. Understand the structure of early Paleozoic extinction events. Understand the major Paleozoic extinction drivers. Understand the degree to which these correlate with Paleozoic extinction events. Early Paleozoic World, Life & Extinctions

Presentation Topics

Stratigraphy - chronostrati- graphy & geochronology Geography - tectonics & distri- bution Climate - circulation, tempera- ture, weather Biota - protists, invertebrates, vertebrates, plants Evolution - evolutionary faunas, adaptive radiations, major innovations Significant Events - sea-level changes, volcanic eruptions, bolide impacts, extinctions Paleozoic World Paleozoic Life

Figure from Fan et al. (2020) Paleozoic World Paleozoic Timescale

Data from ICS (2019) Paleozoic World Phanerozoic Era Durations 300

240

180

120 Duration(myr)

60

0 Paleozoic Mesozoic Cenozoic

Data from ICS (2020) Paleozoic World Paleozoic System Durations

70

56

42

28 Duration(myr)

14

0 Camb. Ord. Sil. Dev. Carbon. Perm.

Data from ICS (2020) Paleozoic World Paleozoic Paleogeography

Animation from Algol (2016: https://www.youtube.com/watch?v=ovT90wYrVk4) Paleozoic World Paleozoic Paleoenvironment

Atmospheric O Atmospheric CO 2 2 35 5000

28 4000

21 3000

14 2000 PercentbyVol. 7 PerMillionParts 1000

0 0 Camb. Ord. Sil. Dev. Carbon. Perm. Camb. Ord. Sil. Dev. Carbon. Perm.

Mean Surface Temperature Sea Level 25 250

20 200

15 150

10 Present 100 Meters Above Meters

DegreesCelsius 5 50

0 0 Camb. Ord. Sil. Dev. Carbon. Perm. Camb. Ord. Sil. Dev. Carbon. Perm. Cambrian Timescale

System/ Numerical Period Series/Epoch /Age Age (Ma) 485.4 ± 1.9 Stage 10 489.5 494.0 Paibian 497.0 500.5 504.5 509.0 Stage 4

Cambrian Series 2 514.0 Stage 3 521.0 Stage 2 529.0 541.0 ± 1.0

ICS International Chronostrat. Chart 2020/03 Cambrian Paleozoic System Durations

70

56

42

28 Duration(myr)

14

0 Camb. Ord. Sil. Dev. Carbon. Perm.

Data from ICS (2020) Cambrian Tectonic Configuration

Laurentia (North America)

Baltica (Scandinavia, eastern Europe, northwestern Russia)

Siberia (Siberia Kazakistan)

Single supercontinent (Gondwana) with three island continents

Map from Scotese PaleoMap Project (2000) Cambrian Marine Circulation

Unimpeded circum-Arctic cold current Broken circum-equatorial current Impeded circum-Antarctic cold current Upwelling zone off southeastern Gondwana

Complex circulation patterns

Map from Scotese PaleoMap Project (2000) Cambrian Paleozoic Paleoenvironment

Atmospheric O Atmospheric CO 2 2 35 5000

28 4000

21 3000

14 2000 PercentbyVol. 7 PerMillionParts 1000

0 0 Camb. Ord. Sil. Dev. Carbon. Perm. Camb. Ord. Sil. Dev. Carbon. Perm.

Mean Surface Temperature Sea Level 25 250

20 200

15 150

10 Present 100 Meters Above Meters

DegreesCelsius 5 50

0 0 Camb. Ord. Sil. Dev. Carbon. Perm. Camb. Ord. Sil. Dev. Carbon. Perm. Cambrian Climate Zones

Comparative Criteria Cambrian

O2 Content of 12.5% vol. % Atmosphere (+63%)

CO2 Content of 4,500 ppm Atmosphere (x16)

Mean Surface 21°C Temperature (+7°C)

Sea Level +30m - 90m

Tropical (Greenhouse) World Broad equatorial arid region High continentality Ice at poles in Early Cambrian, retreating later

Map from Scotese PaleoMap Project (2000) Cambrian Biodiversity

800

Diversification of Cambrian Biota 600

400

Modern Fauna

200 Paleozoic Fauna NumberFamiliesof

Cambrian Fauna 0 Cambrian Sil. Carbon. Tri. Jurassic Cretaceous Tertiary 500 400 300 200 100 0 Geological Time Data from Sepkoski (1981) Cambrian Biodiversity

Figure from Fan et al. (2020) Cambrian Cambrian Evolutionary Fauna

Trilobite Graptolite

Monoplacophoran Inarticulata Hyolith Cambrian Paleozoic Evolutionary Fauna

Articulata Crinoid

Tabulate Coral Ostracod Rugose Coral Cambrian Modern Evolutionary Fauna

Bivalve Gastropod Echinoid Bony Fish Cambrian Cambrian “Explosion”

Increase in O2 levels Increase in calcium concentration in seawater Development of homeobox (Hox) genes, a level of genetic organization that facilities the assembly of different structures in different parts of the body. Initiation of ecological “arms races” “”: sudden Vacant ecospace after appearance of most phyla in extinction the fossil record Complexity threshold breach Cambrian

Cambrian “Explosion” Fauna

Marrella Hallucigenia (Arthropod Acestor) (??? Acestor)

Wiwaxia Archaeopirapula (Mollusc Acestor) (Pirapulid Ancestor) (Crustacean Ancestor) Cambrian Cambrian “Explosion” Fauna

Anomalicaris (Arthropod Ancestor) Cambrian Cambrian “Explosion”

Diagram from Morris (2000) Cambrian Cambrian “Explosion”

Diagram from Cooper & Fortey (1998) Cambrian Reefs

Archaeocyathid Reef Archaeocyathid Reefs The earliest known reefs Constructed by archaeocyathids and receptaculid sponges Archaeocyths are an extinct group of shallow marine, warm-water, sponge- like marine invertebrates especially common in Lower Cambrian sediments.

Map from Scotese PaleoMap Project (2000) Diagram from McKerrow (1978) Cambrian Terrestrial Environments

Naked Land Surface No plants or No soils, but much regolith Complete range of non-biotic surface processes in operation Cambrian Global Climate (Early Cambrian)

Diagram from Hearing et al. (2018) Cambrian Extinctions End-Ordovician End-Devonian End-Permian End-Triassic End-Cretaceous 80 Palaeozoic Mesozoic Cenozoic Cambrian 60

40 PercentExtinction 20

0 Cambrian Ord. Sil. Dev. Carb. Perm. Trias. Jurassic Cretaceous Paleoc. Neo. Paleozoic Mesozoic Cenozoic

Data from Sepkoski (1998) Cambrian Biomere Extinctions

Biomere - Cambrian stage-level, extinction-bounded, trilobite-based biostratigraphic successions characteristic of shallow-water Laurentian sediments in North America.

Figures from Thomas (1995) Cambrian Biomere Extinctions

Allison (Pete) Palmer James Stitt (b. 1927) (19?? - 2016) Cambrian Biomere Extinctions

Tables from Taylor (2006) Cambrian Biomere Extinctions

Interval 4 (Extinction Interval): characterized by Interval 3 survivors.

Interval 3 (Diversification Interval): characterized by endemic speciation + recruitment to achieve further taxonomic + morphological diversification

Interval 2 (Diversification Interval): characterized by endemic speciation + recruitment to achieve enhanced taxonomic diversity.

Interval 1 (Post-extinction Interval) - characterized by recruitment of new species from offshore habitats.

Figure from Taylor (2006) Cambrian Biomere Extinction Causes: Palmer-Stitt Model

Falling sea-level forces continental shelf faunas to migrate to the shelf edges where they compete for limited resources with themselves and shelf-margin species

Eustatic sea-level rise causes the thermocline to migrate onto the continental shelves cooling the marine environment and reducing O2 content.

Diagram from Palmer (1984) Cambrian Biomere Extinction Causes: Saltzman-Knoll Model

Correlatable carbon isotopic excursions in basins from different continents indicate that the pterocephaliid and ptychaspid extinctions were caused by exunic seawater rising onto the continental shelves in a global marine event caused by eustatic sea- level rise.

Steptoean Positive Carbon Isotope Excursion (SPICE)

Most likely both the Palmer-Stitt and Saltzman-Knoll mechanisms were responsible for aspects of the biomere extinctions

Diagram from Saltzman et al (2000) Ordovician Paleozoic World Paleozoic System Durations

70

56

42

28 Duration(myr)

14

0 Camb. Ord. Sil. Dev. Carbon. Perm.

Data from ICS (2020) Ordovician Timescale

System/ Numerical Period Series/Epoch Stage/Age Age (Ma) 443.8 ± 1.5 Hirnantian 445.2 ± 1.4 Upper Katian 453.0 ± 0.7 Sandbian 458.4 ± 0.9 Darriwillian Middle 467.3 ± 1.1 Dapingian

Ordovician 470.0 ± 1.4 Floian Lower 477.7 ± 1.4 485.4 ± 1.9

ICS International Chronostrat. Chart 2020/03 Ordovician

Paleogeography

Gondwana drifts south and rest over the South Pole by the end of the Ordovician Sea-level regression causes Laurentia, Siberia & Baltica to emerge. Massive glaciation in Late Ordovician

Expansion of the Paleo-Tethys ocean w/ rotation of Gondwana, Laurentia, Siberia & Baltica

Map from Scotese PaleoMap Project (2000) Ordovician

Marine Circulation

Unimpeded circum- Arctic cold current Impeded circum- equatorial current No circus-antarctic cold circulation Upwelling zones off western coasts of Laurentia and northern Gondwana

Complex circulation patterns

Map from Scotese PaleoMap Project (2000) Ordovician

Paleozoic Paleoenvironment

Atmospheric O Atmospheric CO 2 2 35 5000

28 4000

21 3000

14 2000 PercentbyVol. 7 PerMillionParts 1000

0 0 Camb. Ord. Sil. Dev. Carbon. Perm. Camb. Ord. Sil. Dev. Carbon. Perm.

Mean Surface Temperature Sea Level 25 250

20 200

15 150

10 Present 100 Meters Above Meters

DegreesCelsius 5 50

0 0 Camb. Ord. Sil. Dev. Carbon. Perm. Camb. Ord. Sil. Dev. Carbon. Perm. Ordovician

Climate Zones

Comparative Criteria Ordovician

O2 Content of 13.5% vol. % Atmosphere (+68%)

CO2 Content of 4,200 ppm (x15) Atmosphere

Mean Surface 16°C (+2°C) Temperature

Sea Level +180m-220m

Temperate ➔ Cold (Icehouse) World Narrowed equatorial arid region High continentality Distinctly expanded southern cold zone at southern pole

Map from Scotese PaleoMap Project (2000) Ordovician

Marine (Neritic) Environment

Great Ordovician Biodiversification Early Ordovician Cambrian- Event (GOBE) fauna acme Diversification of Paleozoic Evolutionary Fauna Archaeocyathid reefs replaced by rugose-tabulate coral & bryozoan reefs. Rise of echinoderms, mollucs and articulate . Replacement of Cambrian with Ordovician descendants. First fish.

Ordovician Marine Scene Ordovician

Terrestrial Environment

Initial invasion of terrestrial habitats First terrestrial plants appear in marginal aquatic environments First arthropods on land as evidenced by preserved trackways Evidence of first soils Evidence of first terrestrial burrows (probably made by arthropods) Away from watercourses, however, the Ordovician landscape was barren

Ordovician Terrestrial Scene Ordovician

Biodiversity

800 Great Ordovician Biodiversification Event (GOBE)

600 End-Ordovician Extinction Event

400

Modern Fauna

200 Paleozoic Fauna NumberFamiliesof

Cambrian Fauna 0 Cambrian Ordovician Sil. Devonian Carbon. Permian Tri. Jurassic Cretaceous Tertiary 500 400 300 200 100 0 Geological Time Data from Sepkoski (1981) Ordovician

Biodiversity

Figure from Fan et al. (2020) Ordovician

Plankton Revolution

Plankton - aquatic species unable to swim against a current (Graptoloidea, Dacryoconarida, Homoctenida, Orthocerids, ).

Demersal Taxa - acquatic species that live in the water column but close to the sea floor (Cephalopoda: , , , Ellesmerocerida, , Lituitida, ; Arthropoda: Radiodonta, Eurypterida; Vertebrata: Galeaspida, Osteostraci, Pteraspidomorphi, Thelodonti; Cephalochordata).

Nekton - aquatic species able to swim and move independent of water currents (, Gnathostomata, most coiled ). Ordovician

Plankton Revolution

Data from Servais et al. (2015) Ordovician

Plankton Revolution

Data from Sepkoski (2002) Data from Paleobiology Database (2014) Ordovician

Great Ordovician Biodiversification Event (GOBE)

Data from Servais & Harper (2018) Ordovician Cambrian Evolutionary Fauna

Trilobite Graptolite

Inarticulata Polychaeta Eocrinoid Ordovician

Paleozoic Evolutionary Fauna

Articulata Crinoid

Cephalopod

Tabulate Coral Bryozoan Rugose Coral Ordovician Modern Evolutionary Fauna

Bivalve Gastropod Echinoid Bony Fish Ordovician

Reefs

Ordovician Reef

First Coral Reefs Note asymmetrical distribution about equator - indicating cold conditions in the Antarctic Ordovician reefs were much more topologically prominent, taxonomically diverrse, and morphologically diverse than their Cambrian counterparts

Map from Scotese PaleoMap Project (2000) Early Paleozoic World, Life & Extinctions Norman MacLeod School of Earth Sciences & Engineering, Nanjing University