CHAPTER 15—The Late Paleozoic World

After completing this chapter, students should:

1. Know the major forms of marine life that declined or disappeared in the late
Paleozoic and the forms that arose to take their place.
2. Be able to describe what happened to reefs after the destruction of tabulate-
strome reefs at the end of the Devonian Period.
3. Understand the ecology of plants of the Carboniferous swamps.
4. Know what kinds of plants lived on higher ground during the Carboniferous.
5. Be able to describe the animals of late Paleozoic freshwater habitats.
6. Know how insects evolved during late Paleozoic time.
7. Be able to describe the rise of reptiles and therapsids and to explain their
evolutionary significance.
8. Know how the positions of the continents shifted during the late Paleozoic.
9. Know the major changes in climate that occurred during the Carboniferous
Period and how these changes affected sea level, sedimentation, and animal
life.
10. Understand the factors that contributed to the distinctness of Permian floras.
11. Be able to explain the interrelationships among atmospheric carbon dioxide,
climate, and plant life in the late Paleozoic.
12. Be able to describe the Late Permian mass extinctions and their probable
causes.
13. Understand why and how the Alleghenian orogeny occurred.
14. Know the late Paleozoic orogenies that occurred in the southwestern United
States.
15. Understand what cyclothems are and how and why they originated in Late
Carboniferous time.
16. Be able to describe the Permian system of west Texas, including its
sedimentation patterns and the evolution of reefs in the Delaware Basin.
17. Know the major tectonic events and sedimentation patterns of the western
margin of North America during late Paleozoic time.

Lecture Outline
I. Overview
A. The late Paleozoic includes the Carboniferous and Permian periods.
B. The late Paleozoic is marked by major climatic changes.
i. Glaciers advanced in Gondwanaland Carboniferous, recedes in
Permian.
ii. Drying of Permian climates caused contraction of coal swamps and
expansion of evaporates.
iii. These changes had major effects on flora and fauna.
C. Two major extinctions of which the Permian Period was the largest ever
experienced
D. Major tectonic change with suturing of Gondwanaland and Euramerica
E. Period names

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 i. The Carboniferous Period was formally recognized in Britain in 1822
to recognize coal-bearing rocks.
1. Americans divided the period in the United States into the
Mississippian, on the basis of limestone in the upper
Mississippi River Valley, and the Pennsylvanian, on the basis
of the coal measures in that state. They are equivalent to Lower
and Upper Carboniferous.
ii. The Permian System was named by Roderick Murchison in 1841 for
rocks in the town of Perm in the Urals of Russia.
II. Late Paleozoic Life—No remarkable differences from that of the Late Devonian
A. New forms of life emerged in Paleozoic seas.
i. Following the extinction of the Late Devonian, there was a
1. Loss of tabulate corals and stromatoporoid sponges
2. Diversification of ammonoids
3. Diverse group of mobile predators, including sharks and ray-
finned fish
4. Loss of Placoderms and nautiloids
a. Loss perhaps due to lack of mobility (too heavily
armored)
5. Rebounding of brachiopods with a new group, the productids,
which contributed to reef development
6. Crinoids reached their greatest diversity during the Early
Carboniferous.
a. Contributed to the Mississippian limestone
b. Bryozoans also contributed to these deposits.
7. Fusulinids (p. 359)—Unusually large single-celled creatures
a. Outstanding index fossils for Upper Carboniferous and
Permian strata
b. Contributed to limestone formation
ii. Aragonitic reef builders flourished in aragonite seas.
1. Coral strome reefs were built by calcite-secreting organisms in
calcite seas.
a. Seas with low magnesium-calcium ratios
2. These seas gave way to aragonite seas and creatures that
secreted aragonite.
a. Aragonite algae in the Late Carboniferous
b. Aragonite sponges in the Permian reefs
iii. Trees grew in swamps.
1. Areas covered by vast wetlands far greater than experienced
today.
2. Small number of genera made up flora.
a. Lepidodendron (p. 360)—A spore-bearing lycopod that
grew to 30 meters height and one meter diameter
b. Sigillaria (p. 360)—Also a spore-bearing lycopod, but
not as large

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 c. New type of fern, seed ferns (p. 360), which used seeds
in addition to spores for reproduction
i. Most abundant was Glossopteris (p. 360), which
is closely associated with Gondwanaland.
iv. Upland floras expanded.
1. Seed ferns and sphenopsids (p. 361) were very abundant on
higher ground. Similar in structure to their modern
descendents, the horsetails
2. Cordaites (p. 361) were another Late Carboniferous group of
tall plants (30 meters) that formed large “pine” woodlands.
a. They are members of the gymnosperm (p. 362) or
naked seed group which forms the modern conifers (p.
362).
3. During the Permian Period, gymnosperms dominated the
terrestrial environment and prevailed during the Mesozoic
Period.
v. Animals diversified on land and invaded freshwater habitats.
1. In freshwater habitats, ray-finned fish continued to diversify
and were joined by freshwater sharks. Also, many species of
bivalves were now found in freshwater environments.
2. Insects—Although they arose in the Devonian, insects
diversified into many species of winged forms during the
Carboniferous.
a. Although the Carboniferous forms could not fold their
wings back, many were of large size. Their presence
indicates higher levels of oxygen.
b. Just before the Permian Period, foldable-wing insects
arose and that group radiated during the Permian
Period.
3. Amphibians
a. Difficult to distinguish between aquatic and terrestrial
amphibians, as they lived in the interface between the
water and land
b. Carboniferous amphibians were very different from
modern ones.
i. Larger in size (some up to six meters)
ii. Terrestrial ones had scales.
4. The rise of the reptiles
a. Oldest known are from Upper Carboniferous
b. Minor differences in skeleton.
c. Major difference in reproduction. Reptiles possessed
amniote egg (p. 364).
i. This egg has four extra-embryonic membranes,
including the amnion.
ii. Allowed the reptiles to lay their eggs away from
water

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 d. They developed a more advanced jaw and teeth that
allowed slicing, in addition to puncturing.
e. Amphibians prospered into early Permian, but during
most of the Permian Period, the reptile began to
displace and replace them.
i. One such group were the pelycosaurs (p. 365),
which included
1. Dimetrodon (p. 365), a fin-backed
reptile the size of a jaguar that had sharp
serrated teeth and could eat small chunks
2. Eryops (p. 365), an alligator-like
amphibian, could only swallow prey
whole.
5. A new level of metabolism
a. Dimetrodon had a structure not unlike the mammals
that evolved from them.
b. Their immediate descendants, the therapsids (p. 365),
had legs positioned more vertically beneath bodies and
complex jaws with differentiated teeth, with large
lateral fangs and bladelike molars.
c. These animals were endothermic (p. 366), which
means they maintained their body temperature by
internal means. Also, suggests they were more active
than other reptiles
d. The other animals, other reptiles and amphibians, were
ectothermic (p. 369), because they maintained their
body temperatures by external means.
III. Paleogeography of the Late Paleozoic World
A. Warm, moist conditions were widespread to Early Carboniferous time.
i. Sea level rose at start. Warm shallow seas spread across the
continental surfaces at low latitudes. Thus, limestones accumulated
over large areas.
ii. The northeastern margin of Euramerica experienced warm, moist
conditions favorable to coal-swamps, whereas the western edge was in
the rain shadow and experienced hot, dry conditions, evaporates and
limestones accumulated in the shallow sea.
B. In mid-Carboniferous time, continents collided and a great ice age began.
i. Gondwanaland collided with Euramerica to produce a supercontinent
called Pangaea.
1. Mountains formed in southern Europe and North Africa.
a. Orogenic activity, therefore, is called the Hercynian
orogeny (p. 369)
2. Mountains formed in North America are the Appalachians
(where the old Acadian orogeny took place) and Ouachita
(Oklahoma and Texas). The orogeny is called the Alleghenian.
ii. Mid-Carboniferous cooling

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 1. Perhaps, because all the carbon was tied up in buried organic
material
2. Began the greatest ice age of the Phanerozoic Eon
a. Lowering of sea level
b. Decimation of marine creatures
c. Major unconformity between Lower and Upper
Carboniferous
d. Steep temperature gradient between poles and equator.
Glaciers moved within 30° of equator.
e. There were cool swamps equatorward of
Gondwanaland and Euramerica, but their flora differed.
iii. Dry habitats expanded in Permian time.
1. Suturing of Siberia to Eastern Europe along the Ural
Mountains completed the assembly of Pangea. Only part
missing was southeast Asia, which attached in the early
Mesozoic.
2. Large size of Pangaea resulted in very acrid conditions mid-
continent. Also, more compartmentalization of the flora.
3. Reduced burial of carbon slowly began to increase the carbon
dioxide levels in the atmosphere, which in turn led to warming
and cessation of glaciation.
4. Two areas did continue to produce coal in the Permian Period.
a. Part of China
b. The Glossopteris growing regions of Pangaea
iv. Mass extinctions ended the Paleozoic Era
1. Two extinctions during the Permian.
a. The first decimated land animals, particularly most of
the therapsids.
b. The second struck most marine creatures and did away
with the rugose and tabulate corals.
IV. Regional Events of the Late Paleozoic Time
A. The Alleghenian orogeny formed the Appalachian Mountains.
i. Mid-Carboniferous collision of Euramerica and Gondwanaland
produced the Alleghenian orogeny (p. 372).
ii. Valley and Ridge Province is zone of low temperature deformation
iii. Eastward was the Piedmont Province: a zone of metamorphosed and
highly deformed rocks because of proximity to the suturing event.
iv. Two zones separated by the Blue Ridge Mountains, a band of
Proterozoic rocks metamorphosed during the Grenville orogeny.
v. Alleghenian orogeny followed quickly on the heels of the Acadian
orogeny. It continued to build new mountains that shed sediments
westward as a molasses.
vi. Modern Appalachian Mountains are a recent isostatic uplift in the area
of this ancient orogenic belt.
B. Orogenies also occurred in the southwestern United States.

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 i. Ouachita Mountains of Oklahoma and Texas are a southwestern
extension of the late Paleozoic Appalachians, but there is a bend and
the connection between the two is buried. There may have been more
than one plate involved. Some parts of the system eventually became
part of Central America.
ii. On the North American craton, areas in the southwestern United States
were transformed into a series of uplifts and basins.
1. Basins accumulated Late Carboniferous and Permian materials,
producing clastic deposits known as arkoses.
a. Fountain Arkose formed along the eastern flank.
b. Today it stands out as a series of spectacular ridges.
iii. The Ancestral Rockies of the Late Carboniferous were produced by
two uplifts, Front Range and Uncompahgre.
1. Perhaps stood as high as the modern Rockies
2. Lay near the equator, so winds produced a rain shadow to the
west, where great thickness of evaporite formed in the Paradox
Basin
C. Coal deposits formed within cyclothems.
i. Western slope of Appalachians consisted of braided streams that
produced a molasses. Most coal swamps were connected at one time or
the other, but there were some separate basins.
1. Michigan
2. Illinois
3. Allegheny
ii. Flatness of the land meant that slight changes in sea level could either
produce freshwater coal swamps or shallow seas. This alternation of
deposition produced the cyclothems (p. 380) of North America and
their equivalent in Britain, the coal measures (p. 380).
iii. Shifting environments and the origin of cyclothems
1. As mentioned above, the western slope of the Appalachians
was almost flat.
2. Therefore, transgressive and regressive seas produced
alternating deposits of marine peats, marine sediments, and
nonmarine sediments.
iv. Glaciers and sea level
1. The above-mentioned changes may have been caused by
repeated expansion and contractions of Gondwanaland
glaciers.
2. Why hasn’t that happened in the Quaternary Period?
a. Perhaps, because the seas are rising and falling over
steeply sloping continental margins rather than the flat
margin of the Carboniferous Period
D. Reefs formed in the Delaware Basin of western Texas.
i. Delaware Basin is located in Texas and New Mexico.
1. Important because of scenery
2. Important because of petroleum reserves

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 ii. It still remains a depositional basin with mountains at the edge that
were once reefs during the Permian Period.
iii. Two basins were formed in the early and Middle Paleozoic, the
Delaware and the Midland basins.
iv. Reef growth
1. During Permian Period, the Midland Basin filled in with
sediments, but the Delaware Basin remained flooded.
2. As sea level rose, the Delaware Basin
a. When shallow, had snails, bivalves, sponges, and
brachiopods
b. With deepening, only the floating creatures ended up in
the sediments: radiolarians, conodonts, and ammonoids.
c. With increased deepening, floor became poorly
oxygenated and few bottom dwellers could survive.
3. The reeflike structure rimming the basin was formed not quite
at the very end of the Permian Period (Guadalupian Age).
a. Formed of aragonitic sponges, algae, and lacy
bryozoans
b. There was an extensive back reef flat.
c. Rubble from the foreslope tumbled into the basin as a
talus slope. It preserved many fossil remains in silica.
d. Basin opened into the ocean toward the southwest via
the Hovey Channel.
e. At the start, the Hovey Channel supplied oxygen-rich
water to the basin bottom, but toward the end, the
channel remained shallow while the basin deepened.
Above low-oxygen conditions resulted.
v. Death of the reef
1. At the end of the Permian Period, the Delaware Basin filled
with evaporate.
2. Hovey Channel may have become restricted.
3. Reef stopped growing. Layering of evaporate may reflect
seasonal changes.
4. Evaporite protected the reef structures, which were later
revealed when freshwater dissolved them.
E. The Sonoma orogeny expanded the North American continent.
i. During late Paleozoic, the western shoreline passed through
northwestern Nevada.
ii. Seaward of the margin was a region of volcanic activity that shed
clastic materials in the basin between arc and shoreline.
iii. Sonoma orogeny was similar to the earlier Antler orogeny.
1. Marine sediments were thrust up on the continental margin.
2. Some were welded onto the continental margin along with the
volcanic terrane of the arc.
3. Complete closure of area between the arc and the continental
margin

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 iv. The above resulted in the westward growth of the North American
crustal plate.

Answers to Textbook Review Questions

1. The expansion and contraction of glaciers in Gondwanaland caused shallow
seas to expand and contract over the midcontinental United States during
Late Carboniferous time.
2. You could determine whether coal deposits formed along a river or a shallow
sea by examining the deposit cycle. River deposits grade upward from coarse
to fine; delta and marine deposits grade upward from fine to coarse.
3. Lycopods, ferns, sphenopsids, and gymnosperms (conifers) still exist in the
modern world.
4. It is only the Early Carboniferous that harbors vast quantities of coal; the Late
Carboniferous is characterized by an unusually large percentage of limestone.
Thus, the limestone-rich Lower Carboniferous is referred to as Mississippian
because of its excellent exposure along the upper Mississippi Valley, and the
coal-rich Upper Carboniferous is referred to as the Pennsylvanian, because of
its widespread occurrence in Pennsylvania.
5. Cycles of falling and rising sea level corresponded to cycles of glacial
expansion and contraction during the Late Carboniferous. As sea level rose
and fell, it created cyclothems of coal deposition. A transgression resulted in
the deposition of marginal marine peat on top of nonmarine deposits; a
regression buried marine deposits beneath peat and then nonmarine
sediments.
6. Several islands collided with the eastern margin of Laurentia during early
Paleozoic time, producing the Taconic orogeny, the first of three Paleozoic
mountain-building episodes that formed the Appalachians. In the middle
Paleozoic, the Acadian orogeny (the second Appalachian mountain-building
episode) resulted from two collisions. First, Laurentia was sutured to Baltica;
later, orogenic activity progressed southward, as Avalonia collided with
eastern Euramerica. The Late Carboniferous collision of Euramerica with
Gondwanaland shifted the region of mountain building along the eastern
margin of North America southward . The fold-and-thrust belt of the central
and southern Appalachians formed at this time in the Alleghenian orogeny.
Then, the Ouachita Mountains were thrust up as a westward continuation of
the Appalachians. The craton to the north and west of the Ouachita Mountains
also underwent tectonic movements; enormous areas in what is now the
southwestern United States became transformed into a series of uplifts and
basins. Two of the uplifts created the Ancestral Rocky Mountains. During the
uplift of the Ouachita range and other uplands, a small fault block rose up
within the west Texas basin, dividing it into the Delaware Basin and the
Midland Basin.
7. Therapsids' legs were positioned more vertically beneath their bodies than the
legs of reptiles or pelycosaurs, making them more mobile. The jaws of
therapsids were more complex, and the teeth of many species were
differentiated into incisors, fangs, and molars. It is thought that the
therapsids were endothermic, which is advantageous because it allows
animals to maintain a sustained level of activity.
8. The Late Permian climate was hot and dry; as the seas receded, they left
behind extensive evaporite sediments.

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9. Extensive burial of reduced carbon in Carboniferous coal swamps markedly
reduced the concentration of carbon dioxide in the atmosphere, weakening
greenhouse warming of the Earth's surface. When climates became drier in
Late Carboniferous and Permian time, levels of atmospheric carbon dioxide
rose as result of reduced rates of weathering and the drying up of coal
swamps (so that less carbon was buried). The increase in atmospheric carbon
dioxide strengthened greenhouse warming.
10. The deep ocean was well oxidized until Guadalupian time. The onset of anoxic
conditions in the deep sea coincided with the first Permian extinction; the
terminal Permian extinction occurred at the time of maximum anoxia of the
oceans. There is evidence that the deep ocean began to stagnate, and anoxic
sediments accumulated in shallower waters. It is possible that brief episodes
of mixing in the ocean carried large quantities of carbon dioxide to surface
waters and the atmosphere, poisoning many forms of life.