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Unraveling mass extinctions: Permian to Early Jurassic onshore-offshore trends of marine stenolaemate bryozoans
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Unraveling mass extinctions: Permian to Early Jurassic onshore-offshore trends of marine stenolaemate bryozoans
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UNRAVELING MASS EXTINCTIONS: PERMIAN TO EARLY JURASSIC
ONSHORE-OFFSHORE TRENDS OF MARINE STENOLAEMATE
BRYOZOANS
by
Catherine Marie Powers
A Dissertation Presented to the
FACULTY OF THE GRADUATE SCHOOL
UNIVERSITY OF SOUTHERN CALIFORNIA
In Partial Fulfillment of the
Requirements for the Degree
DOCTOR OF PHILOSOPHY
(GEOLOGICAL SCIENCES)
December 2009
Copyright 2009 Catherine Marie Powers
ii
ACKNOWLEDGEMENTS
This research was supported by grants from the Geological Society of America, The
Paleontological Society, the American Museum of Natural History, the Yale Peabody
Museum, the USC Department of Earth Sciences, and a USC WiSE grant to my advisor
David J. Bottjer.
I thank Nicole Bonuso, Matthew Clapham, Steve Dornbos, Margaret Fraiser,
Katherine Marenco, Pedro Marenco, and Sara Pruss, my paleo lab mates at USC, for their
friendship and support. My research also benefited greatly from discussions with all of
them. Matthew Clapham and Nicole Bonuso provided assistance in the field in Europe.
I am truly grateful to my advisor David J. Bottjer, for his willingness to take me on
as a student, his encouragement throughout the years, and his seemingly infinite patience.
Thanks to Frank Corsetti for his willingness, no matter how busy he was, to thoroughly
edit papers, proposals, and cover letters, as well as patiently sit through countless hours
of practice talks! I also thank my other committee members Al Fischer, Ken Johnson, and
Suzanne Edmands.
Fellow bryozoologists Paul Taylor, Marcus Key, Joseph Pachut, and Scott Lidgard
provided invaluable insight and made me feel like my research actually mattered. For
that, I am forever grateful.
I want to thank my family for their constant support of my academic pursuits.
Finally, I am very grateful to my husband, Peter, for his love and encouragement, for his
assistance as camp cook and manager, for lugging pounds after pounds of bryozoan-laden
Permian rocks, and for all of the other help he’s provided with this dissertation.
iii
TABLE OF CONTENTS
ACKNOWLEDGEMENTS ii
LIST OF TABLES vi
LIST OF FIGURES vii
ABSTRACT ix
CHAPTER I: INTRODUCTION 1
1. Introduction: Permian to Jurassic 1
2. Mid-Phanerozoic mass extinctions 5
2.1. The end-Guadalupian extinction 5
2.2. The end-Permian extinction and the Early Triassic recovery 6
2.3. The end-Triassic extinction and the Early Jurassic recovery 10
3. Marine stenolaemate bryozoans 11
3.1. Phanerozoic Diversity 11
3.2. Permian-Jurassic diversity 12
3.3. Bryozoan morphology 12
4. Objectives and research implications 16
CHAPTER II: BRYOZOAN PALEOECOLOGY INDICATES 18
MID-PHANEROZOIC EXTINCTIONS WERE THE PRODUCT
OF LONG-TERM ENVIRONMENTAL STRESS
Abstract 18
1. Introduction 18
2. Previous work 20
3. Methods 21
4. Permian to Early Jurassic paleoenvironmental distribution 26
5. Long-term environmental stress 26
6. Implications for the end-Permian and end-Triassic mass extinctions 29
7. Conclusions 31
CHAPTER III: THE EFFECTS OF MID-PHANEROZOIC 33
ENVIRONMENTAL STRESS ON BRYOZOAN DIVERSITY,
PALEOECOLOGY, AND PALEOGEOGRAPHY
Abstract 33
1. Introduction 34
2. Previous work 37
3. Sources of environmental stress in the Late Permian and Late Triassic 40
4. Methods 41
5. Bryozoan generic diversity and extinction rates 42
iv
6. Paleoenvironmental distribution 46
6.1. Bryozoan genera 46
6.2. Bryozoan orders 49
7. Paleogeographic extinction and recovery patterns 54
8. Evolutionary and ecological impact of long-term environmental 56
stress on bryozoan evolution
9. Late Permian terrestrial-marine link 61
10. Conclusions 63
CHAPTER IV: PERMIAN-EARLY JURASSIC TRENDS IN BRYOZOAN 64
COLONIAL MORPHOLOGY
Abstract 64
1. Introduction 65
2. Previous work 66
3. Methods 67
4. Relative abundance and taxonomic distribution of bryozoan 68
growth forms
5. Paleoenvironmental distribution of bryozoan growth forms 72
6. Effects of Late Permian and Late Triassic environmental stress 78
on growth form trends
7. Conclusions 82
CHAPTER V: BEHAVIOR OF LOPHOPHORATES DURING THE 83
END-PERMIAN MASS EXTINCTION AND RECOVERY
Abstract 83
1. Introduction 84
2. Lophophorates 85
3. Early Triassic stenolaemate bryozoans 86
4. Early Triassic brachiopods 88
4.1. Permian survivors 90
4.2. Rhynchonelliform brachiopods 90
4.3. Lingulid brachiopods 91
5. Recovery patterns, timing, and tempo 94
6. Biological and ecological attributes for survival 98
7. Conclusions 101
CHAPTER VI: CONCLUSIONS 104
1. Summary 104
2. Extinction-related environmental stress 105
3. Impact on bryozoan evolution 106
4. Bryozoan colonial growth forms 107
5. Early Triassic lophophorates 108
v
BIBLIOGRAPHY 110
APPENDICES
Appendix A: Permian to Early Jurassic formations 126
Appendix B: Bryozoan references 143
Appendix C: Stratigraphic, environmental, and generic diversity data 190
Appendix D: Bryozoan stage-level generic diversity 205
Appendix E: Bryozoan colonial morphology 221
vi
LIST OF TABLES
Table 2.1: Sedimentological criteria used to assess depositional environments 22
Table 3.1: Permian-Jurassic diversity and extinction rates of stenolaemate 45
bryozoans
Table 3.2: Diversity and extinction rates of stenolaemate bryozoans across 62
environments for the end-Guadalupian and end-Permian extinctions
vii
LIST OF FIGURES
Figure 1.1: Phanerozoic diversity of the three Evolutionary Faunas 2
Figure 1.2: Phanerozoic diversity of marine bryozoan families 3
Figure 1.3: Evidence of protracted Late Permian environmental stability 7
Figure 1.4: Early Triassic recovery patterns 9
Figure 1.5: Triassic diversity of the major bryozoan orders 13
Figure 1.6: Bryozoan colony growth morphologies 15
Figure 2.1: Permian-Early Jurassic bryozoan paleoenvironmental history 23
Figure 2.2: Localities of Early Triassic bryozoan faunas 28
Figure 3.1: Permian-Triassic Paleozoic and Modern Faunas generic diversity 36
Figure 3.2: Permian-Jurassic range of stenolaemate and gymnolaemate bryozoans 38
Figure 3.3: Ordovician-Recent bryozoan generic diversity 43
Figure 3.4: Paleoenvironmental history of Permian-Early Jurassic bryozoans 47
Figure 3.5: Permian paleoenvironmental history of Order Fenestrata 50
Figure 3.6: Permian-Triassic paleoenvironmental history of Order Trepostomata 51
Figure 3.7: Permian-Triassic paleoenvironmental history of Order 52
Cryptostomata and Order Cystoporata
Figure 3.8: Permian-Early Jurassic paleoenvironmental history of Order 53
Cyclostomata
Figure 3.9: Permian-Jurassic paleogeographic distribution of bryozoans 55
Figure 3.10: Jurassic-Tertiary paleoenvironmental history of Order 58
Cheilostomata
Figure 3.11: Phanerozoic paleolatitudinal distribution of bryozoan-rich deposits 59
Figure 4.1: Permian-Early Jurassic relative abundance of bryozoan growth forms 69
viii
Figure 4.2: Permian-Jurassic growth form summary for each bryozoan order 71
Figure 4.3: Permian-Early Jurassic paleoenvironmental history of erect bryozoans 73
Figure 4.4: Permian-Early Jurassic paleoenvironmental history of massive 75
bryozoans
Figure 4.5: Permian-Early Jurassic paleoenvironmental history of encrusting 76
bryozoans
Figure 4.6: Permian-Early Jurassic relative abundance of bryozoan growth 77
forms across environments
Figure 4.7: Phanerozoic relative abundance of the erect and encrusting 79
bryozoans
Figure 5.1: Permian-Triassic range of stenolaemate bryozoans and 87
rhynchonelliform brachiopods orders
Figure 5.2: Paleobiogeographic distribution of Early Triassic bryozoan 89
assemblages
Figure 5.3: Early Triassic paleobiogeographic distribution of rhynchonelliform 92
brachiopod faunas
Figure 5.4: Early Induan paleobiogeographic distribution of lingulid 93
brachiopod assemblages
Figure 5.5: Early Triassic range of rhynchonelliform brachiopods, lingulids, 95
and stenolaemate bryozoans
ix
ABSTRACT
Changes in the diversity and environmental distribution of marine stenolaemate
bryozoans through the Permian to Early Jurassic interval are used to constrain the timing
of environmental stress, determine the degree to which it differentially affected marine
settings, evaluate its long-term impact on bryozoan evolution, and review current
extinction scenarios for the end-Permian and end-Triassic crises. Results indicate that the
end-Permian and end-Triassic mass extinctions were characterized by protracted intervals
of environmental stress that initiated during the end-Guadalupian event several million
years prior to the end-Permian, and again during the Norian stage prior to the end-
Triassic. Environmental degradation lasted into the Early Triassic and Early Jurassic,
hampering the recovery of marine communities. Prior to both extinctions offshore
settings were affected first, suggesting environmental stress resulted from the gradual
encroachment of a deep-water phenomenon onto the shelves, thus precluding catastrophic
events as kill mechanisms and supporting long-term oceanographic processes such as
widespread euxinia resulting from massive volcanism and global warming. The end-
Permian event was the most influential for bryozoans, triggering a permanent change in
their paleoenvironmental preferences from onshore to mid-shelf settings and a taxonomic
switch between stenolaemate and gymnolaemate bryozoans. The taxonomic switch itself
initiated a gradual shift in the overall morphological composition of bryozoan
assemblages between erect-dominated Paleozoic communities and encruster-dominated
Mesozoic communities. Major changes in the paleoenvironmental preferences of
bryozoans were also coupled with fluctuations in their diversity. Elevated bryozoan
x
extinction rates were concurrent with their disappearance from offshore settings in the
Late Permian and Late Triassic. Re-colonization of vacated habitats took place as
diversity slowly rebounded in the wake of the extinctions, indicating a return to normal
marine conditions. A comparison of survival patterns of stenolaemate bryozoans with
other lophophorates during the end-Permian extinction revealed that bryozoans were the
most susceptible, experiencing the highest rates of extinction and becoming
geographically restricted during the Early Triassic. The most successful lophophorate
group, the lingulid brachiopods, possessed a unique physiological adaptation that allowed
them to survive in environmentally stressed Early Triassic settings.
1
CHAPTER I
Introduction
1. Introduction: Permian to Jurassic
The Permian to Jurassic interval is a pivotal transition in the evolution of life,
marked by three large biotic crises, the end-Guadalupian, end-Permian and end-Triassic
extinctions, and the most significant marine faunal reorganization of the Phanerozoic
(Raup and Sepkoski, 1982; Sepkoski, 1981; Stanley and Yang, 1994). Members of the
“Modern Fauna”, mainly bivalves and gastropods, successfully weathered the mass
extinctions to become the dominant constituents of marine benthic communities,
replacing the brachiopod/crinoid-dominated “Paleozoic Fauna” (Fig. 1.1) (Sepkoski,
1981, 1984). The ecological and biological consequences of these extinctions (the end-
Permian in particular) have been the subject of intense paleontological research, largely
focusing on the “Paleozoic to Modern Fauna” transition, with particular emphasis on the
nature of the bivalve takeover from brachiopods of level bottom benthic marine
environments in the Early Triassic (e.g., Erwin, 2001; Fraiser and Bottjer, 2007b; Gould
and Calloway, 1980; Jablonski, 2001; Sepkoski, 1984).
Concurrently, a significant but less well-known taxonomic turnover occurred
within the Phylum Bryozoa; whereas stenolaemate bryozoans dominated Paleozoic
communities, gymnolaemates were the dominant post-Paleozoic group (Fig. 1.2) (Taylor
and Larwood, 1990). Although only one order of bryozoans became extinct at the
Permian-Triassic boundary, the remaining groups exhibited a delayed recovery and low
2
Figure 1.1. Phanerozoic diversity trends of the three Evolutionary Fauna.
The transition between the Paleozoic and Modern Fauna coincides with
the end-Permian mass extinction. Arrows indicate times of major mass
extinction discussed in this study (1 = end-Guadalupian, 2 = end-Permian,
3 = end-Triassic). Modified from Sepkoski (1981).
3
Figure 1.2. Bryozoan family diversity curve. Shaded area indicates the
time interval for the proposed study. Modified from Taylor and Larwood
(1988).
4
diversity throughout the Triassic (Powers and Pachut, 2008; Schäfer and Fois, 1987).
Before they could fully recover, stenolaemate bryozoans nearly went extinct at the end of
the Triassic; only one order survived to later diversify in the Jurassic (Powers and Pachut,
2008; Taylor and Larwood, 1988). Pre-extinction diversity levels were not re-established
until the mid-Cretaceous, ~150 m.y. later (Fig. 1.2), with the advent of a new
gymnolaemate order Cheilostomata, but bryozoans never regained their widespread
Paleozoic distribution. These taxonomic changes have been documented but questions
concerning the impact of the end-Guadalupian, end-Permian and end-Triassic extinctions
remain. What environmental and ecological processes shaped bryozoan evolution in the
early Mesozoic? Specifically, how did the environmental distribution of bryozoans
change across the Permian-Triassic and Triassic-Jurassic boundaries and were these
changes gradual or abrupt? Finally, what can these changes in their environmental
distribution tell us about the extent of the environmental degradation during the Late
Permian and end-Triassic extinctions? This dissertation attempts to answer all of these
questions and to provide a comprehensive account of the taxonomic and ecological
effects of three successive mass extinctions on one of the major invertebrate phyla of the
Paleozoic. I maintain that the constant, and sometimes dominant, presence of bryozoans
in marine benthic communities throughout the Phanerozoic and their ability to adapt to
changing environmental conditions make them a suitable and useful proxy to examine the
environmental effects of mass extinctions on marine communities.
5
2. Mid-Phanerozoic mass extinctions
2.1. The end-Guadalupian extinction
The end-Guadalupian extinction (259 Ma, Zhou et al., 2002) is a Late Permian
event, first identified by Raup and Sepkoski (1982) who combined it with the end-
Permian extinction as a single crisis, and then acknowledged as an independent extinction
by Stanley and Yang (1994). Until recently, extinction estimates ranged from 58 to 65%
for skeletonized marine genera (Knoll et al., 1996; Stanley and Yang, 1994), but a recent
study incorporating published data of newly described marine fossils from the Late
Permian revealed that the end-Guadalupian extinction may not have been as severe, with
a total generic extinction rate of only 28.1% for marine invertebrates (Clapham and
Bottjer, 2007a). Most affected by the end-Guadalupian were ammonoids, trilobites, and
bryozoans, all with generic extinction rate above 40% (Clapham and Bottjer, 2007a). The
end-Guadalupian extinction has also been recognized as a distinct crisis in the terrestrial
realm, with generic extinction among vertebrates at about 67% (Retallack et al., 2006).
Although still a matter of contention, volcanism (Emeishan igneous province),
global sea level fall, and a late Middle Permian high bio-productivity period (the
“Kamura” event) are all potential kill mechanisms (Hallam and Wignall, 1999; Isozaki et
al., 2007; Zhou et al., 2002). However, recent geochemical and lithologic data suggest
that the causal mechanism was gradual, which precludes catastrophic processes (i.e.,
extraterrestrial impact, gas hydrate release), and that the development of deep-water
anoxia associated with the P/T superanoxia event was initiated at the end of the Middle
6
Permian suggesting that extinction-related environmental instability was likely protracted
throughout the Late Permian (Fig. 1.3) (Isozaki, 1997; Isozaki et al., 2007).
2.2. The end-Permian extinction and the Early Triassic recovery
The end-Permian mass extinction (252 Ma, Mundil et al., 2004), one of the most
devastating biotic crises of the Phanerozoic, is marked by the disappearance of about
80% of marine species and 49% and 63% of marine and terrestrial families, respectively
(Benton, 1995; Raup and Sepkoski, 1982; Stanley and Yang, 1994). Whereas sessile,
filter-feeding organisms (i.e., bryozoans, brachiopods, corals, and echinoderms) were
affected most by the extinction, mobile organisms such as gastropods and bivalves
proliferated in the subsequent aftermath (Fraiser and Bottjer, 2004, 2007b; Knoll et al.,
1996; Knoll et al., 2007). Extinction in the terrestrial realm was contemporaneous with
high diversity drops recorded for insects, amphibians, and reptiles (Benton, 1988, 1989,
1995; Hallam and Wignall, 1997).
Hypothetical causal mechanisms for the end-Permian extinction include a range
of gradual and catastrophic processes: widespread oceanic and atmospheric anoxia (Huey
and Ward, 2005; Isozaki, 1997; Wignall and Twitchett, 1996), hypercapnia (Knoll et al.,
1996), euxinia (H
2
S poisoning) (Grice et al., 2005; Kump et al., 2005; Nielsen and Shen,
2004), massive volcanism and global warming (Kamo et al., 2003; Renne et al., 1995),
methane oxidation (Krull and Retallack, 2000; Ryskin, 2003), and a meteorite impact
(Becker et al., 2001). The fundamental kill mechanism remains unclear and is likely a
combination of several of the aforementioned processes (Erwin, 2006). Whereas evidence
7
Figure 1.3. Protracted environmental stability during the late Permian. A
= Negative long-term trend in the δ
13
C
carb
values across a Capitanian-
Wuchiapigian section in Japan. Modified from Isozaki et al. (2007). B =
Permian-Triassic superanoxia event showing the development of deep-
water anoxic conditions at the end of the Guadalupian (Wuch. –
Wuchiapingian, Chang. – Changsingian, Olenek. – Olenekian). Modified
from Isozaki (1997).
8
for a catastrophic extraterrestrial scenario is not compelling (Farley et al., 2005; Isozaki,
2001), geochemical, sedimentological, and paleontological data suggest that the end-
Permian crisis was the result of a prolonged interval of environmental degradation caused
by widespread euxinia triggered by massive volcanism and global warming (Clapham
and Bottjer, 2007b; Grice et al., 2005; Powers and Bottjer, 2007; i.e., Renne et al., 1995;
Ward, 2006).
The subsequent marine recovery in the Early Triassic was delayed by several
million years. Metazoan reefs of any kind were absent for most of the Early Triassic and
many groups of organisms did not fully recover until the Middle Triassic (Fig. 1.4) (see
Hallam and Wignall, 1997 and references therein; Stanley, 1988). Early Triassic marine
invertebrate faunas were depauperate and dominated by a few abundant and
cosmopolitan species of ammonoids, bivalves, gastropods, and lingulid brachiopods
(Fraiser and Bottjer, 2004, 2005; Rodland and Bottjer, 2001; Schubert and Bottjer, 1995).
Sustained environmental instability throughout the Early Triassic, documented by
unusual sedimentary features (i.e., large sea-floor carbonate precipitates, flat-pebble
conglomerates, wrinkle structures, and microbial buildups in normal marine settings) and
repeated large-scale disturbances in the carbon isotopic record, likely contributed to the
low marine diversity and delay in recovery (Fig. 1.4) (i.e., Baud et al., 1996; Lehrmann et
al., 2001; Payne et al., 2004; Pruss et al., 2006; Pruss et al., 2004; Schubert and Bottjer,
1995; Woods et al., 1999).
9
Figure 1.4. Early Triassic recovery patterns. A = Faunal patterns of common marine invertebrates across the Permian-
Triassic boundary. Modified from Hallam and Wignall (1997). B = Unusual sedimentary features and carbon isotopic
record during the Early Triassic (M – Middle; E – Early). Modified from Pruss et al. (2006).
10
2.3. The end-Triassic extinction and the Early Jurassic recovery
The end-Triassic extinction (201 Ma, Shaltegger et al., 2008), another of Raup
and Sepkoski’s (Raup and Sepkoski, 1982) five largest mass extinctions, is marked by the
disappearance of 80% of marine invertebrate species (Sepkoski, 1996), including many
ammonite families, bivalve genera, and reef organisms (i.e., scleractinians, sponges)
(Beauvais, 1984; Hallam, 1981; Tozer, 1981). In contrast, marine vertebrates were not
significantly affected (Benton, 1991; Hallam and Wignall, 1997).
Postulated extinction mechanisms include CO
2
and SO
2
emissions associated with
flood basalt volcanism of the Central Atlantic Magmatic Province (CAMP), gas hydrate
release from seafloor sediments, and a meteorite impact (Beerling and Berner, 2002;
Cohen and Coe, 2002; Hesselbo et al., 2002; Olsen et al., 2002; Pálfy et al., 2001). Like
the end-Permian extinction, the exact kill mechanism is still unknown, although Ward
(2006) suggested that it may have been analogous to that of the end-Permian and
triggered by increased CO
2
emissions in the atmosphere and widespread oceanic euxinia.
Anoxic conditions have only been reported from several Late Triassic sections in Europe
and British Columbia (Ciarapica, 2007; Sephton et al., 2002; Van de Schootbrugge et al.,
2007; Ward et al., 2004), which indicate it may have only been a regional phenomenon
(Tanner et al., 2004).
Similar to the end-Permian aftermath, reefs exhibited a delayed recovery (~ 8 my,
Pliensbachian Stage, Fraser et al., 2004), but most other marine invertebrate faunas
rapidly reestablished themselves (within 1 my) during the first stage of the Jurassic
(Hallam and Wignall, 1997). However, the late Early Jurassic was marked by another
11
small extinction event immediately above the Pliensbachian-Toarcian boundary that is
attributed to widespread anoxia resulting from rapid sea level rise (Aberhan and
Baumiller, 2003; Hallam, 1986; Jenkins, 1988). Organisms affected by this event include
bivalves, rhynchonellid brachiopods, and ostracods (Aberhan and Baumiller, 2003;
Hallam, 1986; Whatley, 1988).
3. Marine stenolaemate bryozoans
3.1. Phanerozoic diversity
The Phanerozoic diversity of bryozoan families reveals two important radiations
during their evolutionary history (Fig. 1.2). An initial radiation took place in the
Ordovician after their first appearance in marine communities and was followed by a
relatively stable plateau through the remainder of the Paleozoic, with relatively minor
losses at the end-Ordovician and Devonian extinctions. The second radiation did not
initiate until the middle Jurassic; bryozoan diversity has continued rising ever since. This
post-Paleozoic radiation is of critical interest here as it is associated with the end-
Guadalupian and end-Permian mass extinctions and the subsequent events of the Triassic.
Superimposed over these diversity changes are changes in the taxonomic composition of
bryozoan communities. Paleozoic bryozoans were dominated by the Orders of the Class
Stenolaemata while post-Triassic communities were, and still are today, dominated by
representatives of the Class Gymnolaemata (Taylor and Larwood, 1988). Although the
latter evolved in the Ordovician, their contribution to Paleozoic bryozoan diversity was
negligible.
12
3.2. Permian-Jurassic diversity
Bryozoan diversity dropped dramatically at the Permian-Triassic boundary (Fig.
1.5). The decline of bryozoans in the Late Permian started at the end of the Guadalupian
with an estimated 52-61% generic extinction (Gilmour and Morozova, 1999; Knoll et al.,
1996; Stanley and Yang, 1994; Taylor and Larwood, 1988); the end-Permian itself is
responsible for an estimated 64-69% generic extinction (Stanley and Yang, 1994; Taylor
and Larwood, 1988). At the ordinal level, only one order (of six), the Fenestrida, became
extinct. The remaining orders exhibited a delayed recovery and low diversity throughout
the Triassic (Fig. 1.5) (Powers and Pachut, 2008; Schäfer and Fois, 1987). The recovery
of bryozoans was not complete until the late Middle Triassic when bryozoans rapidly
diversified. Analyses of paleogeographic distribution indicate that the Tethys Ocean was
a locus of origination and a center of bryozoan diversity from which bryozoans
eventually extended their ranges into higher latitudes (Powers and Pachut, 2008).
Stenolaemate bryozoans nearly went extinct at the end of the Triassic; only one order
survived to later diversify in the Jurassic (Powers and Pachut, 2008; Taylor and Ernst,
2008; Taylor and Larwood, 1988).
3.3. Bryozoan Morphology
Bryozoans are colonial animals composed of thousands of genetically identical
individuals (zooids) that secrete a calcium carbonate skeleton ranging in mineralogy from
low-magnesium calcite to aragonite (Cuffey and Utgaard, 1999; Schopf and Manheim,
1967; Taylor, 1999). Each zooid harbors a bryozoan individual and has a specific
13
Figure 1.5. Triassic bryozoan diversity. A = Permian to Early Jurassic
bryozoan generic diversity. E: Early, M: Middle, L: Late, J: Jurassic. Early
Jurassic data is from Taylor (pers. commun.). B = Triassic bryozoan
species diversity subdivided into the four dominant orders. Trias.: Triassic,
In.: Induan, Olen.: Olenekian, Rha.: Rhaetian. Modified from Powers and
Pachut (2008).
14
function within the colony. Bryozoan colonies can range in size from microscopic, in
which they are composed of only a few zooids, up to 1 m (Boardman and Cheetham,
1987; McKinney and Jackson, 1989). The addition and arrangement of zooids control
colony growth and morphology (Taylor, 1999). Four basic colony growth forms are
associated with bryozoans. These growth forms include encrusting, erect, massive, and
free living (Fig. 1.6). Encrusting colonies can grow as single or multilayered sheets of
zooids. Erect colonies are attached at the base to hard substrates, grow upward through
the addition of zooids, and can consist of uni-, bi-, or multiserial sheets or branches. The
degree of calcification determines the rigidity of the colony and its flexibility (Boardman
and Cheetham, 1987). Erect colonies may be branching with the branches connecting in
some cases to form a net-like pattern (Boardman and Cheetham, 1987; McKinney and
Jackson, 1989). Free-living colonies are unattached and generally have a discoid shape
(Boardman and Cheetham, 1987).
Whereas internal processes (i.e., colonial integration, communication between
zooids) control colonial growth form, the distribution of these forms is often a result of
extrinsic factors (Hageman et al., 1997 and references within; McKinney and Jackson,
1989). These generally include such environmental and ecological variables as
bathymetry, substrate, water energy, temperature, food availability, and competition.
Research has shown that changes in the frequency of colonial morphotypes may reflect
changes in the environment. In fact, one of the most striking patterns is the switch from
the Paleozoic erect-dominated to the post-Paleozoic encruster-dominated faunas
15
Figure 1.6. Bryozoan colony growth morphologies. Free-living colony is
not shown. Modified from Hageman et al. (1998).
16
(McKinney and Jackson, 1989), which may be connected to the environmental and biotic
crises of the mid-Phanerozoic.
4. Objectives and research implications
The primary goals of this project are to 1) use marine stenolaemate bryozoans as a
proxy for assessing extinction-related environmental change during the Permian to
Jurassic interval, and 2) evaluate the long-term evolutionary impact of the end-
Guadalupian, end-Permian, and end-Triassic mass extinctions on bryozoan evolution. To
address both of these questions, datasets of the diversity, distribution (global and
environmental), and morphology of marine bryozoans were collected and plotted. Time-
environment diagrams served as the main tool for investigating the environmental health
of marine bryozoan communities during the Late Permian and end-Triassic extinctions
and are therefore emphasized throughout the dissertation.
The data and results presented have important implications for mass extinction
research and our current understanding of bryozoan evolution. Extinction intensity in
different marine environments characterized by bryozoan environmental trends can be
linked to proposed mass extinction mechanisms and used to assess the validity of each.
Any future scenario about the possible cause of a mass extinction must now account for
disparity in environmental degradation along marine shelves. The environmental impact
of the Late Permian and end-Triassic extinctions also now provides a context for the
major evolutionary changes that took place within Phylum Bryozoa during the early
Mesozoic.
17
Many of the results and implications of the research presented in this dissertation
are now published in peer-reviewed journals. Chapter II, published in Geology (Powers
and Bottjer, 2007), focuses on the severity of mass extinctions in varying environments
during the Permian to Early Jurassic interval and its implications for mass extinction kill
mechanisms. Chapter III, published in Global and Planetary Change (Powers and
Bottjer, 2009), investigates the long-term impact of the Late Permian and end-Triassic
mass extinctions and their associated environmental degradation on the diversity,
distribution, and evolution of marine stenolaemate bryozoans. Chapter V, published in
the Journal of Asian Earth Sciences (Powers and Bottjer, 2008), surveys how specific
adaptations to changing environmental conditions may have helped certain marine
invertebrate groups survive in the aftermath of the End-Permian mass extinction.
18
CHAPTER II
Bryozoan paleoecology indicates mid-Phanerozoic extinctions were the
product of long-term environmental stress
Abstract
I compiled the global onshore-offshore distribution of marine bryozoans within
432 Permian to Early Jurassic marine assemblages and show that bryozoan assemblage
generic richness declined significantly in advance of the end-Permian and end-Triassic
mass extinctions, starting as early as the Capitanian prior to the end-Permian and the
Norian prior to the end-Triassic. I also show that offshore settings were affected first,
prior to both extinctions, suggesting environmental stress resulted from the gradual
encroachment of some deep-water phenomenon onto the shelves. These patterns support
long-term oceanographic, rather than extraterrestrial, extinction mechanisms such as
widespread euxinia triggered by massive volcanism and global warming. Tracking the
onshore-offshore environmental distribution of these marine invertebrates provides a
unique approach to assessing prolonged environmentally induced stress through this ~120
m.y. time interval.
1. Introduction
The end-Permian and end-Triassic mass extinctions are considered to be two of
the largest biotic crises of the Phanerozoic. A range of mechanisms, from oceanographic
to climatic to extraterrestrial, has been proposed to explain both events (e.g., Becker et
al., 2001; Grice et al., 2005; Isozaki, 1997; Knoll et al., 1996; Krull and Retallack, 2000;
19
Olsen et al., 2002; Renne et al., 1995; Tanner et al., 2004; Wignall and Twitchett, 1996),
but the fundamental kill mechanism(s) remains unclear. Traditionally, a major drop in
standing diversity generally defines mass extinctions, as displayed by compilations of
taxonomic richness (Sepkoski, 1981), and the environmental context of the taxa in
question is not always considered. How does mass extinction-related environmental
stress differentially affect the marine settings (i.e., nearshore, offshore, reefs) in which
the organisms that eventually became extinct were living? Because the severity of
environmental stress on marine environments immediately prior to extinctions is
undeniably linked to causal mechanisms, it must be assessed before considering
extinction models. In this study, I compiled the global onshore-offshore distribution of
marine stenolaemate bryozoans through time to assess the environmental context of these
extinctions. I then applied these data to evaluate current extinction scenarios for both
events.
Marine bryozoans have a long and rich fossil record from level-bottom and reefal
communities dating back to the Ordovician Period (Boardman and Cheetham, 1987).
Their Phanerozoic diversity parallels the trends documented by Sepkoski (1981) for all
marine invertebrates. Detailed studies on Permian to Jurassic bryozoans have focused on
their diversity and paleogeographic distribution (e.g., Gilmour and Morozova, 1999;
Morozova, 1969; Powers and Pachut, 2008; Schäfer and Fois, 1987; Taylor and Ernst,
2008), and provide a framework for understanding how they were affected by both the
end-Permian and end-Triassic extinctions as well as their survival and recovery in the
aftermath of each crisis. Marine bryozoans are also sensitive indicators of environmental
20
conditions; their diversity, species distribution, and morphology have been found to
correlate with various environmental factors (e.g., Amini et al., 2004; Hageman et al.,
1997; Taylor, 2005). Tracking the environmental distribution of bryozoans therefore
enables us to test the severity of mass extinctions in a range of settings.
2. Previous work
Previous onshore-offshore studies have successfully been used to track the
environmental histories of major groups of organisms and have revealed several large-
scale Phanerozoic trends such as patterns of originations of higher taxa (Jablonski and
Bottjer, 1991; Jablonski et al., 1983) and evolutionary novelties (Jablonski et al., 1997),
the expansion of members of the three Evolutionary Faunas during the Ordovician
radiation (Sepkoski and Miller, 1985; Sepkoski and Sheehan, 1983), the environmental
distribution of post-Paleozoic benthic marine invertebrates (Bottjer and Jablonski, 1988),
Phanerozoic extinction trends across environmental settings (Sepkoski, 1987), bivalve
environmental trends in the Paleozoic (Miller, 1988), and Phanerozoic trends in the
environmental distribution of trace fossils (Bottjer et al., 1988).
Only two studies have dealt directly with bryozoans. A study by Jablonski et al.
(Jablonski et al., 1997) analyzed the environmental origination of evolutionary novelties
characteristic of cheilostome and cyclostome bryozoans during the post-Paleozoic. An
earlier and more pertinent study by Bottjer and Jablonski (1988) tracked the
environmental history of cheilostome bryozoans from their origination in the Late
Jurassic to the Tertiary. Bottjer and Jablonski (1988) observed a trend toward increasing
21
generic diversity in offshore settings coupled with a rapid diversification in the Late
Cretaceous. Cheilostomes also became dominant in middle and outer shelf environments
following the end-Cretaceous mass extinction. While this study focused on
gymnolaemate bryozoans, it provides significant evidence that long-term evolutionary
trends of higher taxa can be tracked and may be affected by large environmental
disturbances such as mass extinctions.
3. Methods
I compiled a global data set of 432 marine fossil stenolaemate bryozoan assemblages
from the Permian through the Jurassic (Appendix A), culled from primary literature,
taxonomic monographs, and the Paleobiology and PaleoReef Databases (Appendix B).
Bryozoan assemblages were assigned to seven broad environmental zones along an
onshore-offshore gradient: 1) nearshore settings above fairweather wave base, including
high-energy settings such as shoals and bars; 2) inner shelf and lagoon, above normal
storm wave base; 3) middle shelf environments between normal and maximum storm
wave base; 4) reefs and bioherms; 5) outer shelf, immediately landward of the shelf edge;
6) deep-water slope mud mounds; and 7) slope and basin settings beyond the shelf edge.
Each depositional setting was defined based on physical processes, mainly water energy,
as reflected in the type of sediments and its grain size, fossil preservation, and
sedimentary features (e.g., Bottjer and Jablonski, 1988) (Table 2.1). The most generically
diverse bryozoan faunal assemblages for each environmental category were then plotted
against time to create a time-environment (T-E) diagram (Fig. 2.1). Detailed generic
22
Environmental bin Sedimentary characteristics
Nearshore 1. Subtidal but above fairweather base; includes shoals and bars.
2. Thick laterally-continuous beds of sand and coarser sediments with parallel laminae and cross-bedding
3. Fossils disarticulated, abraded, well-bedded
Inner shelf & lagoon 1. Above normal storm wave base; includes deltaic settings
2. Common storm beds with parallel laminae or hummocky cross-stratification interbedded with fairweather fine-
grained beds
3. Fossils commonly in storm beds
Middle shelf 1. Between normal and maximum storm wave base
2. Massive fine-grained beds, commonly rhythmic, storm beds relatively uncommon
3. Fossils unabraded
Reefs & bioherms 1. Subtidal
2. Resistant framework, individual mound or lens-shaped buildups, can be laterally extensive
3. In situ frame builders
Outer shelf 1. No storm influence, landward of shelf edge
2. Massive fine-grained strata, commonly rhythmic on a fine scale, no evidence of slumping, mass movement, or
turbidites
3. Fossils unabraded
Slope mounds 1. Downslope of shelf edge
2. Mud-dominated, poorly sorted, lenticular to conical form with steep edges
3. Lack significant in situ frame-building organisms
Slope and deep basin 1. Beyond shelf edge
2. Slumping, mass movement, turbidites, deep-sea fan facies geometries; fine-grained sediments and microbreccias
3. Fossils from beds deposited as turbidites or by some other type of mass movement omitted
Table 2.1. Sedimentological criteria used to assess depositional environments. Modified from Bottjer and Jablonski (1988).
23
Figure 2.1. Contoured Permian-Early Jurassic time-environment (T-E) diagram of
marine bryozoans. Each dot represents a data point, either for the assemblage with the
greatest bryozoan generic richness in each T-E bin, or for absence of bryozoans validated
by the taphonomic control group. Horizontal axis represents environmental zones;
vertical axis represents time in millions of years (Gradstein and Ogg, 2004). Stratigraphic
and locality information and references are available in Appendices B and C. Early
Permian: A=Asselian, S=Sakmarian, Ar=Artinskian, K=Kungurian; Middle Permian
(MP): R=Roadian, W=Wordian, C=Capitanian; Late Permian (LP): W=Wuchiapingian,
C=Changsingian; Early Triassic (ET): *=Induan, O=Olenekian; Middle Triassic:
A=Anisian, L=Ladinian; Late Triassic: C=Carnian, N=Norian, R=Rhaetian; Early
Jurassic: H=Hettangian, S=Sinemurian, P=Pliensbachian, T=Toarcian. Bryozoan
assemblages are composed of the following stenolaemate orders: Permian Fenestrata;
Permian-Triassic Trepostomata, Cryptostomata, and Cystoporata; and Permian-Jurassic
Cyclostomata. Contouring was done separately for each time Period using cubic
interpolation. Four phases of bryozoan paleoenvironmental history are: A, wide
distribution across all environments from Early into Middle Permian, followed by
Middle-Late Permian decline in offshore settings; B, patchy distribution during Early
Triassic; C, gradual re-colonization of marine environments during Middle-early Late
Triassic but restriction to middle shelf and reefal settings in latest Triassic; D, restriction
to shelf settings during Early Jurassic.
24
25
richness for each assemblage used is available in Appendix C. Several assemblages,
particularly in the Triassic and Early Jurassic, only include one genus. The time axis is
divided into 20 intervals ranging from the Early Permian to Early Jurassic, with each time
bin representing a different stage based on the International Stratigraphic Chart
(Gradstein and Ogg, 2004). The resulting matrix contains 140 T-E bins. Fossil
assemblages were assigned to a time bin using conodont, fusulinid, or brachiopod
biostratigraphic data. In instances where no bryozoan genera were reported in a particular
T-E bin, I used a taphonomic control group, the rhynchonelliform Brachiopoda, to
ascertain whether the absence of bryozoans was real or may represent a preservational or
sampling bias. Rhynchonelliform brachiopods and bryozoans are lophotrochozoans that
share a similar skeletal mineralogy and lophophorate filter-feeding life habit, and both are
commonly found together in marine communities. The presence of these brachiopods in
certain environments implies that the absence of bryozoans from those settings is a real
phenomenon, not a taphonomic artifact. Unequal sampling among environmental bins
and differential preservation of bryozoan faunal assemblages caused by facies variation
were accounted for by selecting and plotting the most generically diverse bryozoan
assemblage for each environmental category on the T-E diagram.
This representation of the environmental distribution of marine bryozoans as a
group might possibly lead to an oversimplified interpretation of the dataset by not
considering the different ecological requirements of each stenolaemate order
(Cryptostomata, Cyclostomata, Cystoporata, Fenestrata, and Trepostomata) (e.g., Schopf,
1969). However, an examination of the T-E histories of each group in Chapter II revealed
26
that the individualistic tendencies that characterize each stenolaemate order do not play a
role in determining the paleoenvironmental distribution of bryozoans during times of
significant environmental stress.
4. Permian to Early Jurassic paleoenvironmental distribution
The resulting T-E diagram (Fig. 2.1) displays the contoured maximum generic
richness of bryozoan assemblages. During the Early and Middle Permian, bryozoans
occupied niches across all environmental zones, but through the Middle and Late
Permian steadily declined in offshore deep-water settings. Across the Permian-Triassic
boundary during the Early Triassic, bryozoans show no distinct onshore-offshore trend,
with only 1-2 genera in some environments while being absent in other settings.
Bryozoans once more expanded to colonize most marine environments during the Middle
and early Late Triassic, with their highest generic richness in reefs and mounds. In the
latest Triassic, bryozoans gradually became restricted to middle shelf and reefal settings,
having disappeared from both nearshore and deep-water environments. Early Jurassic
bryozoans were largely concentrated in shelf settings.
5. Long-term environmental stress
The T-E history of marine bryozoans clearly reflects the influence of mass extinction-
related environmental stress on Permian-Early Jurassic marine ecology. In the Permian,
bryozoans declined in deep-water environments first, suggesting that deep-seated
environmental degradation was initiated in the Middle Permian before the end-
27
Guadalupian crisis (ca. 259 Ma). The sporadic environmental distribution and low local
generic richness of bryozoan assemblages during the immediate aftermath of the end-
Permian extinction mirrors their low global diversity and narrow geographic range during
the Early Triassic (seven species belonging to four genera are documented from only four
locations) (Fig. 2.2). The dearth of bryozoans in reefs and slope mounds is a direct result
of the absence of metazoan reefs of any kind through most of the Early Triassic, a
common characteristic of post-extinction intervals throughout the Phanerozoic (Bottjer,
2004). This protracted biotic recovery (~5 m.y., Lehrmann et al., 2006) is symptomatic
of prolonged environmental stress in Early Triassic oceans, as documented by unusual
sedimentary features and repeated large-magnitude carbon isotopic excursions (Payne et
al., 2004; Pruss et al., 2006). The narrow geographic distribution of bryozoan genera in
the Early Triassic also provides strong evidence for prolonged environmental stress.
Bryozoans were restricted to high latitude environments along the western and
northwestern coast of Pangea in the eastern Panthalassic Ocean and, to date, no Early
Triassic bryozoans have been documented from the Tethys Ocean, where the recovery of
typical metazoan reefs eventually took place (Powers and Pachut, 2008) (Fig. 2.2). These
boreal settings may have served as refugia where bryozoans and other marine organisms
could have escaped the vicissitudes of the Early Triassic oceans (e.g., Wignall et al.,
1998).
The increased environmental versatility of bryozoans in the Middle and Late Triassic
mirrors the return of normal marine conditions, faunas, and metazoan reef communities.
However, the disappearance of bryozoans from offshore and nearshore settings, and their
28
Figure 2.2. Localities where Early Triassic bryozoan species have been
described. Locality A: 2 species; Locality B: 1 species; Locality C: 3
species; Locality D: 1 species. Modified from Powers and Pachut (2008).
29
restriction to reefal and middle shelf environments in the latest Triassic once again
highlights the influence of looming mass extinctions on marine communities. Bryozoan
decline in the Late Triassic started during the Norian Stage. The Early Jurassic is
characterized by another delayed recovery of metazoan reefs (Fraser et al., 2004; Stanley,
1988). The global diversity of Early Jurassic bryozoans was low (10 genera) with their
geographic distribution patchy and restricted to the Tethys realm (Taylor and Ernst,
2008).
6. Implications for the end-Permian and end-Triassic mass extinctions
I can use the observed pattern surrounding the Permian-Triassic and Triassic-
Jurassic boundary intervals to evaluate extinction scenarios for both crises. The
protracted nature of bryozoan decline, coupled with the important fact that deep faunas
declined before shallow faunas, cannot be explained by some catastrophic extraterrestrial
impact. Recently, widespread oceanic euxinia (anoxia and hydrogen sulfide poisoning)
triggered by massive volcanism (Siberian traps for the end-Permian and CAMP for the
end-Triassic extinctions) and global warming has been suggested as a kill mechanism
(e.g., Cohen and Coe, 2002; Grice et al., 2005; Hesselbo et al., 2002; Kump et al., 2005;
Mundil et al., 2004; Nielsen and Shen, 2004; Ward, 2006; Wignall and Twitchett, 1996).
The occurrence of deep-water anoxia during the Late Permian (Isozaki, 1997) and
the eventual incursion of euxinic waters into the photic zone at the end of the Permian
(Grice et al., 2005; Riccardi et al., 2007; Wignall et al., 1998) have been demonstrated in
several areas. Such conditions are consistent with fluctuations in atmospheric O
2
and CO
2
30
as hypothesized by recent models (Berner, 2006). As O
2
concentrations in the oceans
dropped, the chemocline separating oxic from euxinic water masses rose to the surface,
bringing along water rich in CO
2
and H
2
S that invaded shallow water environments and
proved to be detrimental to marine life in the photic zone through hypercapnia and
sulphide toxicity (e.g., Grice et al., 2005; Knoll et al., 1996; Kump et al., 2005; Wignall
and Twitchett, 1996). In this scenario, these conditions affected deeper settings first,
killing bryozoan communities in those environments, before gradually invading shallow
shelves and restricting bryozoans onshore (Fig. 2.1). Stagnating oxygen-poor and sulfide-
rich waters along shallow shelves killed those remaining organisms unable to cope
physiologically with the elevated CO
2
and H
2
S content of the water. Evidence of
persisting euxinic conditions during the Early Triassic at several boundary sections (e.g.,
Japan, British Columbia, Meishan in southern China, Spitsbergen, and the Dolomites)
provides additional evidence for the prolonged environmental stress that led to the
delayed recovery of most marine invertebrates and the patchy occurrence of bryozoans in
Early Triassic environments (Grice et al., 2005; Isozaki, 1997; Wignall and Twitchett,
1996, 2002).
Evidence of anoxic or euxinic conditions in the Late Triassic is not as conclusive
(e.g., Tanner et al., 2004). Widespread anoxia has been reported from several sections in
British Columbia and Europe, indicating that anoxic conditions were present in shallow
and deep-water settings as early as the end of the Norian (Ciarapica, 2007; Sephton et al.,
2002; Van de Schootbrugge et al., 2007; Ward et al., 2004). Ocean-atmosphere modeling
of the Late Triassic based on elevated pCO
2
(McElwain et al., 1999) also suggests
31
enhanced oceanic stratification and decreased levels of O
2
in the water column during the
Late Triassic, supporting increased CO
2
as the cause of environmental stress in Late
Triassic oceans (Huynh and Poulsen, 2005).
The disappearance of bryozoans in offshore environments in the Late Triassic
coincides with the onset of anoxic conditions in marine basins, similar to the pattern
observed in the Permian. The similarities between the end-Permian and end-Triassic mass
extinctions (Siberian Traps versus CAMP, increased atmospheric CO
2
, carbon cycle
perturbations) and the decline of bryozoans from deep-water settings concurrent with the
onset of anoxic conditions strongly suggest analogous intrinsic mechanisms for both
extinctions.
7. Conclusions
This study indicates that intrinsic factors, such as long-term oceanographic
mechanisms, were responsible for the shift in habitats exhibited by bryozoans during the
Permian-Triassic and Triassic-Jurassic intervals. The decline of bryozoans in deep-water
settings beginning in the Middle Permian and then again in the Norian indicates that both
extinctions were part of two prolonged intervals of environmental stress caused by the
gradual shallowing of anoxic to euxinic conditions. This contrasts with recent works in
which the end-Permian and end-Triassic mass extinctions have been viewed as
catastrophic events (e.g., Bowring et al., 1998; Olsen et al., 2002; Rampino et al., 2000),
and instead revives the older perception of mass extinctions, and in particular the end-
Permian event, as slow protracted crises (e.g., Teichert, 1990). The gradual nature of the
32
Middle-Late Permian bryozoan decline also implies that the end-Guadalupian and end-
Permian crises were linked by one prolonged episode of environmental instability (e.g.,
Clapham and Bottjer, 2007a) and may not represent two separate bryozoan extinctions.
The differential environmental trends exhibited by nearshore bryozoans at the
Permian-Triassic and Triassic-Jurassic boundaries suggest a potential varied role for
stressful atmospheric and terrestrial perturbations on Late Permian and Late Triassic
shallow marine environments (e.g., Fraiser and Bottjer, 2007a). Other linkages between
climate change and the spatial distribution of marine and terrestrial organisms have been
suggested from both the fossil record and modern realm (Hughes, 2000; Montañez et al.,
2007), implying that environmental factors can exert more control over the range of a
species than ecological dynamics. Tracking the broad environmental patterns of
organisms can thus provide a unique opportunity to examine the impact of
environmentally induced stress into a range of settings.
33
CHAPTER III
The effects of mid-Phanerozoic environmental stress on bryozoan diversity,
paleoecology, and paleogeography
Abstract
Evidence of sustained environmental degradation associated with the end-
Guadalupian, end-Permian, and end-Triassic extinctions has been inferred from
numerous geochemical and sedimentological studies, but the long-term impacts of this
extinction-associated stress on the evolutionary trajectories of marine invertebrates have
not been explored. An examination of the diversity, extinction, paleoenvironmental
range, and geographical distribution of marine stenolaemate bryozoans during the
Permian to Jurassic interval provides striking new evidence of the taxonomic and
ecological influence of these mid-Phanerozoic extinctions on one of the most abundant
components of the Paleozoic Fauna. Elevated bryozoan extinction rates during the Late
Permian and Late Triassic were coupled with major changes in their habitats. Bryozoans
gradually disappeared from deep-water offshore settings during the Late Permian and
from nearshore and offshore settings during the Late Triassic. Re-colonization of these
environments in the wake of each crisis was delayed but coupled with increases in global
generic diversity. The taxonomic effects of the end-Guadalupian extinction were milder
than previously described, even though ecologically, bryozoans were becoming restricted
to nearshore settings. The end-Permian mass extinction remained the largest for
bryozoans, drastically reducing global and assemblage generic diversity and triggering a
permanent change in their paleoenvironmental preferences from nearshore to mid-shelf
34
settings. The 285 million year dominance of stenolaemate bryozoans ended during the
Late Triassic when all but one order (Cyclostomata) became extinct, initiating a
taxonomic switch between stenolaemate and gymnolaemate bryozoans. Moreover, spatio-
temporal variations in the paleoenvironmental history of bryozoans imply that Late
Permian and Late Triassic marine environmental instability resulted largely from some
stressful deep-water phenomenon. High extinction rates in nearshore environments in the
Late Permian provide a link between marine and terrestrial/atmosphere extinction-related
perturbations.
1. Introduction
The transition between Paleozoic and Mesozoic communities is characterized by
three devastating crises (Raup and Sepkoski, 1982). The end-Guadalupian (Middle
Permian) and end-Permian extinctions are two independent events that took place within
8 million years (Myr) of each other in the Late Permian (Gradstein et al., 2004; Mundil et
al., 2004; e.g., Stanley and Yang, 1994). Together, they contributed to the most severe
drop in biodiversity of the Phanerozoic and a major shift in the taxonomic and ecological
structure of marine communities (Clapham and Bottjer, 2007a; Erwin, 2006; Fraiser and
Bottjer, 2005, 2007b; Gould and Calloway, 1980). Fifty-two million years later at the end
of the Triassic, many of the Paleozoic survivors who had struggled to re-diversify in the
wake of the Late Permian events became extinct. These three extinctions, spanning about
61 Myr, were accompanied by at least two periods of sustained environmental instability.
Late Permian environmental degradation likely started at the end of the Middle Permian
35
and persisted through the Early Triassic (e.g., Algeo et al., 2007; Grice et al., 2005; Huey
and Ward, 2005; Isozaki, 1997). Late Triassic environmental stress inferred from ocean-
atmosphere modeling based on elevated pCO
2
(Huynh and Poulsen, 2005; McElwain et
al., 1999) was likely initiated during the Rhaetian (Ciarapica, 2007).
The end-Guadalupian and end-Permian extinctions preferentially terminated
sessile members of the Paleozoic Fauna (i.e., bryozoans, brachiopods, and crinoids)
(Knoll et al., 1996; Knoll et al., 2007). While all groups suffered considerable losses at
the end of the Permian, members of the Modern Fauna (i.e, bivalves, gastropods) fared
better in the immediate aftermath. Knoll et al. (2007) reviewed the effects of
environmental perturbations on both faunas and concluded that extinction selectivity in
the Late Permian was linked to environmental stress, specifically hypercapnia.
Hypercapnia, or CO
2
-poisoning, is one of several proposed mechanisms for the end-
Permian extinction; others include various oceanographic, climatic, and extraterrestrial
processes (Erwin, 2006). Regardless of the trigger, Late Permian-Early Triassic
environmental stress had a more lasting effect on members of the Paleozoic Fauna.
Secular changes in biodiversity during the Permian-Triassic (P/T) interval indicate that
the re-diversification of sessile organisms during the Early Triassic was slow and that
pre-extinction (Middle Permian) diversity levels were not yet re-established as of the Late
Triassic (Fig. 3.1).
In this study, I examined the transition of bryozoans through the Permian to
Jurassic interval in an attempt to constrain the effects of mass extinction-related
environmental stress on their evolutionary history. Stenolaemate bryozoans, diverse and
36
Figure 3.1. Changes in the generic diversity of members of the Paleozoic
and Modern Faunas across the end-Guadalupian and end-Permian
extinctions. Modified from Knoll et al. (1996). Grey area represents the
duration of extinction-related environmental stress based on
sedimentological and geochemical evidence (Isozaki, 1997; Payne et al.,
2004; Pruss et al., 2004; Woods et al., 1999).
37
abundant members of the Paleozoic Fauna, nearly became extinct at the end of the
Triassic (one group, Cyclostomata, survived) and were replaced as the dominant post-
Jurassic group by the Modern Fauna gymnolaemate bryozoans (Fig 3.2) (Sepkoski, 1981;
Taylor and Larwood, 1990). Analyses of bryozoan diversity changes, paleoenvironmental
transitions, and paleogeographic distribution have the potential to reveal why these mid-
Phanerozoic extinctions had a much more significant impact on some groups (i.e, the
Paleozoic Fauna) than others.
2. Previous work
Until recently, diversity and paleoecological studies of marine bryozoans were
largely confined to the Paleozoic (excluding the late Permian) and the post-Triassic due
to the perceived paucity and lack of knowledge of bryozoan taxa in Late Permian to Early
Jurassic assemblages (Taylor and Larwood, 1988). The only detailed field investigation
of bryozoans across a Permian-Triassic boundary interval dates back more than four
decades (Morozova, 1965). Knowledge of Late Permian and Triassic bryozoan diversity
increased rapidly during the 1980s and coincided with more detailed investigations of
Permo-Triassic boundary intervals (i.e., Morozova and Zharnikova, 1984; Sakagami,
1985; Schäfer and Fois, 1987; Schäfer and Fois-Erickson, 1986).
Yet, comprehensive bryozoan diversity and paleoecological patterns during mass
extinction intervals, particularly for the end-Permian and end-Triassic events, have rarely
been quantified and included in studies of mass extinctions and their recoveries, and the
role of environmental disturbances on the evolutionary trajectory of bryozoans has never
38
Figure 3.2. Schematic illustrating the range of stenolaemate and gymnolaemate (Gymno) bryozoans through the
Permian to Jurassic interval. Data from Powers & Pachut (2008).
39
been fully explored. Instead, investigations of other large marine invertebrates (i.e, corals,
brachiopods, ammonoids) and micro-invertebrates (i.e., foraminifera) have taken
precedence, largely due to their biostratigraphic utility (see for example Beauvais, 1984;
Mei and Henderson, 2001; Tozer, 1981; Waterhouse and Bonham-Carter, 1976).
The most recent and comprehensive, but separate, diversity compilations of Late
Permian, Triassic, and Jurassic bryozoans provide a good springboard for further
paleoecological and paleoenvironmental assessment of bryozoans through two of the
most critical evolutionary events of the Phanerozoic. The Middle-Late Permian bryozoan
generic compilation by Gilmour and Morozova (1999) is based on years of field
observations and collections by both authors and certainly represents one of the most
complete diversity and paleogeographic studies of Middle-Late Permian bryozoans.
However, while their compilation provides a good assessment of diversity in different
climatic zones (each of which being further divided into individual provinces based on
Late Permian paleogeography), only partial faunal lists for each of the locations
mentioned in the paper are included. Furthermore, their analysis does not take into
account the paleoenvironmental settings of the collected assemblages. Detailed analyses
of Triassic and Jurassic bryozoan species and generic diversity are provided in Powers
and Pachut (2008) and Taylor and Ernst (2008) but provide no paleoenvironmental
context either.
40
3. Sources of environmental stress in the Late Permian and Late Triassic
Environmental stress in the Late Permian and the Late Triassic is linked to the
end-Guadalupian, end-Permian, and end-Triassic mass extinctions. Negative carbon
isotopic excursions across the end-Guadalupian boundary, a gradual trend towards
decreasing values during the early Late Permian, and lithological evidence of the
development of deep-water anoxia suggest that extinction-related environmental
instability was likely protracted through the Middle-Late Permian boundary and is the
likely source of the sustained Late Permian environmental degradation (Isozaki, 1997;
Isozaki et al., 2007). The P/T superanoxia event is linked to the development of deep-
water anoxia initiated during the end-Guadalupian crisis and, in addition to other
geochemical, sedimentological and paleontological data, indicate that Late Permian
environmental stress lasted through the Early Triassic (Clapham and Bottjer, 2007b;
Isozaki, 1997; Payne et al., 2004; Pruss et al., 2006; Pruss et al., 2004; Woods et al.,
1999).
While evidence of anoxic or euxinic conditions in the late Triassic is not as
conclusive as during the Late Permian-Early Triassic interval (e.g., Tanner et al., 2004),
anoxic conditions have been reported from several Late Triassic sections in Europe and
British Columbia (Ciarapica, 2007; Sephton et al., 2002; Van de Schootbrugge et al.,
2007; Ward et al., 2004) and ocean-atmosphere modeling of the Late Triassic suggests
enhanced oceanic stratification and decreased levels of oxygen
in the water column
during the Late Triassic (Huynh and Poulsen, 2005). Negative carbon isotopic excursions
from several sections around the world also provide compelling evidence for
41
environmental instability during the Triassic-Jurassic (T/J) boundary interval (Hesselbo
et al., 2002; McElwain et al., 1999; Pálfy et al., 2001; Ward et al., 2001).
4. Methods
I created two global data sets using bryozoan marine fossil assemblages in order
to quantify the diversity, extinction, and distribution patterns of bryozoans through the
Permian to Jurassic interval. Data were culled primarily from published literature,
taxonomic monographs, and the Paleobiology Database (www.paleodb.org) (Appendix
B).
In the first compilation, I recorded the presence of marine stenolaemate bryozoan
genera at the stage level to calculate new extinction percentage rates for the end-
Guadalupian, end-Permian, and end-Triassic events (Appendix D). Previously calculated
rates for these extinctions were outdated in light of many recently published taxonomic
studies and improvements in P/T and T/J biostratigraphy that enhanced the time-
stratigraphic resolution of many bryozoan assemblages previously assigned to older
broad time units (i.e., Kazanian, Dzhufian).
The second compilation recorded the generic composition, geographic and
stratigraphic location, and depositional environment of 432 marine bryozoan assemblages
from the Early Permian through the Early Jurassic (Appendix A). This is the same
compilation used in Chapter II and the data were plotted in a similar fashion (see Chapter
II for detailed methodology). However, the dataset was subdivided to construct time-
environment (T-E) diagrams for each of the major stenolaemate bryozoan orders to
42
further determine the degree to which environmental factors controlled the onshore-
offshore distribution of marine bryozoans.
Finally, the paleogeographic distribution of bryozoan faunal assemblages
combined with paleoenvironmental data was plotted on paleogeographic reconstructions
(Scotese, 2001) to examine geographical trends across the end-Guadalupian, end-
Permian, and end-Triassic extinction intervals.
5. Bryozoan generic diversity and extinction rates
The Permian to Jurassic generic diversity of stenolaemate bryozoans is
characterized by an abrupt extinction in the Late Permian and low diversity levels
throughout the Triassic and Jurassic (Fig. 3.3). Pre-extinction diversity levels of marine
bryozoan genera were not re-established until the mid-Cretaceous, ~150 m.y. later (Fig.
3.3). New Late Permian extinction rates, calculated using our genus-level compilation but
excluding stratigraphic singletons, differ from previously published rates for bryozoans,
but fall within the intensity range for all marine invertebrates (Table 3.1). Bryozoan
extinction intensity steadily increased throughout the Late Permian, with extinction of
35.3% (29/82) at the end of the Guadalupian and 73.5% (25/34) for the end-Permian
mass extinction. The end-Triassic mass extinction seemed the most severe, with 75%
(3/4) of bryozoan genera becoming extinct. However, the standing diversity of bryozoan
genera in the latest Triassic had been severely diminished by a relatively high extinction
at the Norian/Rhaetian boundary (60%) (Powers and Pachut, 2008) and was much lower
than at the end of the Permian (Table 3.1), therefore enhancing the magnitude of the end-
43
Figure 3.3. Ordovician-Recent generic diversity. Arrows indicate the end-Guadalupian,
end-Permian, and end-Triassic extinctions. Shaded areas represent the Permian to
Jurassic time interval. EP = Early Permian (Asselian, Sakmarian, Artinskian, Kungurian),
MP = Middle Permian (Roadian, Wordian, Capitanian), LP = Late Permian
(Wuchiapigian, Changsingian), * = Early Triassic (Induan, Olenekian), MT = Middle
Triassic (Anisian, Ladinian), LT = Late Triassic (Carnian, Norian, Rhaetian), EJ = Early
Jurassic (Hettangian, Sinemurian, Pliensbachian, Toarcian), MJ = Middle Jurassic
(Aalenian, Bajocian, Bathonian, Callovian), LJ = Late Jurassic (Oxfordian,
Kimmeridgian, Tithonian). Ordovician to Carboniferous data from Horowitz and Pachut
(2000); Jurassic data from Taylor and Ernst (2008); Early Cretaceous data from Pachut
(pers. comm. 2007); Middle Cretaceous to Holocene data from McKinney and Taylor
(2001).
44
45
Bryozoans Previous Studies %
Diversity Extinction % Bryozoans All Invertebrates
End-
Guadalupian
82 29 35.3 52-61 28.1-59.5
End-Permian 34 25 73.5 28-69 56.1-77.6
End-Triassic 4 3 75 n/a 41-48
Table 3.1. Diversity, extinction, and extinction rate of stenolaemate bryozoan for the end-Guadalupian, end-Permian,
and end-Triassic extinctions. Also displayed for comparison are extinction rates for bryozoans and all invertebrates
from previous studies (Clapham and Bottjer, 2007a; Gilmour and Morozova, 1999; Kiessling et al., 2007; Knoll et al.,
1996; Sepkoski, 1986; Stanley and Yang, 1994; Taylor and Larwood, 1988).
46
Triassic extinction. The increasing rates of extinction throughout the Late Permian and
Late Triassic may be symptomatic of protracted extinction-related environmental stress.
6. Paleoenvironmental distribution
6.1. Bryozoan genera
The time-environmental history of bryozoan assemblages is displayed in Figure
3.4. Several trends are discernable at different time scales, i.e., stage vs. period level. At
the longer time-scale, it seems that bryozoans display their highest diversity in nearshore
environments throughout the Permian and in mid-shelf settings during the Triassic and
Early Jurassic, suggesting that a switch in the environmental preferences of bryozoans
took place during the P/T interval. A closer look reveals nearly three cycles of expansion
and retreat over the entire Permian to Early Jurassic interval (Fig. 3.4). First, an offshore
expansion during the early stages of the Permian was followed by a gradual retreat (or
extinction) from offshore deep-water to nearshore settings during the Middle and Late
Permian. The disappearance of bryozoans from deep-water settings is coincident with the
increase in bryozoan extinction rate throughout the Late Permian. The
paleoenvironmental trend exhibited by Late Permian bryozoans emphasizes the
ecological impact of the end-Guadalupian and end-Permian extinctions.
A second offshore expansion took place during the Middle and early Late Triassic
during which bryozoans extended their range to all marine settings before gradually
retreating to middle shelf settings during the T/J interval. Unlike the P/T interval, no
sudden change in the paleoenvironmental distribution of bryozoans can be seen across the
47
Figure 3.4. Contoured maximum generic diversity of Permian-Early Jurassic marine
stenolaemate bryozoan assemblages. Each dot represents a data point, either for a
bryozoan assemblage or for the absence of bryozoans validated by the taphonomic
control group. Horizontal axis represents environmental zones; vertical axis represents
time in millions of years (Gradstein and Ogg, 2004). Stratigraphic and locality
information and references are available in Appendices B and C. Contouring was done
separately for each time Period using cubic interpolation. E = Early, M = Middle, L =
Late, NS = Nearshore, IS & L = Inner Shelf & Lagoon, MS = Middle Shelf, R = Reefs,
OS = Outer Shelf, SM = Slope Mounds, S & DB = Slope & Deep Basin.
48
49
T/J boundary, further accentuating the difference between both mass extinctions. Finally,
a third, smaller bi-directional expansion, characterized by a slow re-colonization of inner
and outer shelf environments, seems to have taken place in the late Early Jurassic. Each
expansion and retreat is correlated with changes in the global diversity of bryozoans, with
diversity increasing during expansion periods and vice versa (Fig. 3.3). Both retreats took
place immediately prior to the extinction intervals in the Permian and Triassic and are
most likely related to increased environmental stress.
6.2. Bryozoan orders
To determine the actual mechanisms (intrinsic biological processes vs.
environmental factors) driving the environmental distribution of bryozoans through the
Permian to Jurassic interval (Fig. 3.4), I also examined the T-E history of each
stenolaemate bryozoan order (Fenestrata, Trepostomata, Cryptostomata, Cystoporata, and
Cyclostomata) (Figs. 3.5-3.8). The data used to make these diagrams comes from the
same assemblages used in Figure 3.4.
Fenestrates, which became extinct at the end of the Permian, were clearly the
most dominant bryozoan order of the Permian with high diversity in nearly all
environments (Fig. 3.5). They are especially abundant in nearshore and reef settings and
are the only group with high generic richness in deep slope and basin settings. Fenestrates
are missing from nearshore environments at the end of the Middle Permian, which may
be a response to increasing environmental stress leading to the end-Guadalupian
extinction. Trepostomes were consistently more diverse in nearshore and outer shelf
50
Figure 3.5. Contoured maximum generic diversity of Permian marine bryozoan of the Order Fenestrata. See Fig. 3.4
for explanation of abbreviations.
51
Figure 3.6. Contoured maximum generic diversity of Permian-Triassic
marine bryozoan of the Order Trepostomata. See Fig. 3.4 for explanation
of abbreviations.
52
Figure 3.7. Contoured maximum generic diversity of Permian-Triassic marine bryozoan orders. A = Cryptostomata, B
= Cystoporata. See Fig. 3.4 for explanation of abbreviations.
53
Figure 3.8. Contoured maximum generic diversity of Permian-Early
Jurassic marine bryozoan of the Order Cyclostomata. See Fig. 3.4 for
explanation of abbreviations.
54
settings during the Permian but dominated all Triassic settings, particularly during the
Early Triassic (Fig. 3.6). Trepostomes, along with cryptostomes and cystoporates,
became extinct during the Late Triassic. Both cryptostomes and cystoporates seem to
dominate nearshore settings during the Permian but were extremely rare in the Triassic
(Fig. 3.7). Finally, cyclostomes, the only extant stenolaemate order, were very rare in the
Permian and have not yet been reported from Early and Middle Triassic settings.
However, they account for all of the diversity in Early Jurassic environments (Fig. 3.8).
These different T-E patterns imply that distinct clade histories driven by the
individualistic biological tendencies of each stenolaemate order underlie the
paleoenvironmental trends of stenolaemate bryozoans as a group.
7. Paleogeographic extinction and recovery patterns
The paleogeographical distribution of all known marine bryozoan assemblages
was plotted across each extinction interval (Fig. 3.9). The distribution of bryozoans
across the end-Guadalupian and end-Triassic extinctions remained similar across each
extinction interval. However, a striking switch in assemblage distribution took place
across the P/T boundary. Bryozoan assemblages that were widespread during the Late
Permian became restricted to high latitude settings along northwest Pangea in the eastern
Panthalassa. These patterns further emphasize the important ecologic and taxonomic
implications of the end-Permian mass extinction.
55
Figure 3.9. Paleogeographic distribution of all marine stenolaemate
bryozoan assemblages across the end-Guadalupian, end-Permian, and end-
Triassic extinction intervals. Base maps modified from Scotese (2002).
56
8. Evolutionary and ecological impact of long-term environmental stress on
bryozoan evolution
Bryozoan diversity and distribution through the Permian to Jurassic interval
clearly indicate that prolonged Late Permian and Late Triassic environmental stress in
marine environments had a considerable impact on the evolutionary history of
stenolaemate bryozoans. Extinction rates during the Late Permian and Late Triassic were
elevated above background rates and Late Permian originations were suppressed. The
declines in diversity were coupled with gradual changes in the environmental range of
bryozoan genera, suggesting that the effect of extinction-related stress varied among
marine settings and between mass extinctions. However, the different T-E histories of
each stenolaemate order prior to the Late Permian and Late Triassic imply that during
times of reduced stress, intrinsic biological processes largely drove the
paleoenvironmental distribution of bryozoans.
The end-Guadalupian extinction was not as severe as previously described for
bryozoans (Gilmour and Morozova, 1999), but instead can be considered the first in a
series of events that ended the 285 million year dominance of marine stenolaemate
bryozoans. The lack of geographical trends in the distribution of Capitanian and
Wuchiapingian assemblages (Fig. 3.9) indicates that the extinction was global, as shown
by Gilmour and Morozova (1999) in their analysis of Late Permian bryozoan faunal
provinces.
The end-Permian extinction remains the largest, eliminating all but nine
stenolaemate genera belonging to five families. These Paleozoic holdovers, most of them
trepostomes, dominated the Triassic recovery, both in terms of diversity and
57
environmental extent (Fig. 3.6). In addition to the considerable loss in biodiversity, the
environmental distribution of bryozoans through the P/T interval provides striking new
evidence of the influence of extinction-related stress on the marine realm. Late Permian
bryozoan assemblage generic richness declined significantly starting at the end-
Guadalupian boundary and bryozoans did not occupy deep offshore settings. These
patterns imply that environmental degradation resulted from the gradual encroachment of
some deep-water phenomenon onto the shelves. As discussed in section 2, sources of
Late Permian environmental stress are numerous but most notably include the
development of deep-water anoxia initiated at the end of the Middle Permian (Isozaki,
1997).
A second interesting trend to emerge from the spatial distribution of bryozoans is
the apparent long-term switch in their environmental preferences coincident with the end-
Permian mass extinction. The post-Jurassic T-E history of cheilostome bryozoans (see
Bottjer and Jablonski, 1988) (Fig. 3.10) and preliminary data on Middle and Late Jurassic
cyclostomes indicate that assemblage generic richness has indeed remained higher in
middle and outer shelf environments since the beginning of the Mesozoic.
Contemporaneous to this environmental switch is a change in the global latitudinal
distribution of bryozoan-rich deposits from a Paleozoic tropical to a modern non-tropical
distribution (Fig. 3.11) (Taylor and Allison, 1998). Both of these long-term changes in
environmental preferences correspond to the taxonomic turnover between stenolaemate
and gymnolaemate bryozoans that gradually came about during the Late Permian to
Jurassic interval.
58
Figure 3.10. Jurassic to Tertiary paleoenvironmental history of
cheilostome bryozoan generic richness (Pal = Paleoecene, Eoc = Eocene,
Oli = Oligocene). Modified from Bottjer and Jablonski (1988).
59
Figure 3.11. Trend in the paleolatitudinal distribution of bryozoan-rich
carbonate deposits during the Phanerozoic. Modified from Taylor and
Allison (Taylor and Allison, 1998).
60
Geographical trends in bryozoan assemblages across the P/T boundary reveal a
third, albeit short-lived, change in environmental preferences. The restricted occurrences
of Early Triassic bryozoans outside of Tethys compared to their otherwise cosmopolitan
distribution within Tethys in the Late Permian suggests that boreal settings may have
served as refugia for bryozoans. In fact, geochemical, sedimentological, and
paleontological evidence indicate that environmental instability in high latitude settings
was short-lived and the recovery of marine ecosystems there occurred much more rapidly
than in Early Triassic equatorial environments (Pruss and Bottjer, 2004; Wignall et al.,
1998).
Extinction intensity for the end-Triassic extinction seems larger than for the end-
Permian. However, the standing diversity of bryozoans in the Triassic was extremely low
compared to the Permian, especially in the latest Triassic. Extinction was already
elevated following the relative Carnian high in bryozoan diversity (Powers and Pachut,
2008) and at least one of the remaining four stenolaemate orders became extinct before
the end of the Triassic (Fig. 3.2). Two other stenolaemate orders became extinct during
the Norian stage, bringing an end to almost all Paleozoic holdovers and the stenolaemate
285 million year history of dominance. While Late Triassic environmental stress is not
causally related to the prolonged Late Permian-Early Triassic instability, both episodes
contributed to the decline in diversity and assemblage generic richness of bryozoans that
began with the end-Guadalupian extinction, and continued to suppress originations
throughout the Triassic and the early Jurassic (Powers and Pachut, 2008; Taylor and
Ernst, 2008).
61
9. Late Permian terrestrial-marine link
As reviewed in a previous section of this chapter, the gradual and protracted
nature of Late Permian-Early Triassic and Late Triassic environmental degradation was
the result of oceanographic and atmospheric processes (Grice et al., 2005; Huey and
Ward, 2005; see Isozaki, 1997; McElwain et al., 1999), although contention over the
exact nature of the relationship between events in the marine and terrestrial realm still
exists. Environmental trends in bryozoan extinction rates prior to the end-Guadalupian
and end-Permian mass extinction provide further evidence for the link between terrestrial
and marine perturbations. For both events, extinction intensity was highest in nearshore
environments (Table 3.2). A global end-Permian extinction of rooted plants, which
contributed to a significant increase in terrestrial erosion and sediment influx into the
oceans, and a biocalcification crisis resulting from elevated atmospheric CO
2
may have
caused increased instability in coastal settings (Fraiser and Bottjer, 2007a; Sephton et al.,
2005; Ward et al., 2000). While higher extinction rates in nearshore environments are
expected during background times of extinction due to changing and unpredictable
conditions, extinction intensity generally increases offshore during times of mass
extinction (Sepkoski, 1987). If reefs are excluded because their placement along the shelf
can vary significantly, extinction intensity from the inner shelf to the outer shelf does
increase (Table 3.2), thus highlighting the anomalous extinction intensity in nearshore
settings.
62
End-Guadalupian End-Permian
Diversity Extinction % Diversity Extinction %
Nearshore 18 11 61.1 17 9 52.9
Inner Shelf & Lagoon 28 9 32.1 11 4 36.3
Middle Shelf 44 18 40.9 13 6 46.2
Reefs 23 4 17.4 27 12 44.4
Outer Shelf 24 12 50 2 1 50
Slope Mounds 3 1 33.3 0 0 0
Slope & Deep Basin 0 0 0 0 0 0
Table 3.2. Diversity, extinction, and extinction rate of stenolaemate bryozoan across environments for the end-
Guadalupian and end-Permian extinctions.
63
10. Conclusions
Diversity and spatio-temporal analyses of Permian to Jurassic marine
stenolaemate bryozoans reveal that long-term Late Permian and Late Triassic
environmental instability had a major impact on the evolutionary trajectory of bryozoans,
facilitating a taxonomic switch between stenolaemate and gymnolaemate bryozoans and a
long-term ecological change in their environmental and geographical preferences. The
importance of this protracted extinction-related stress on marine settings and organisms
reinforces the notion of an oceanographic process as the trigger of both the Late Permian
and Late Triassic extinctions. Integrating global diversity studies of fossil marine
organisms with their long-term paleoenvironmental trends allows us to better constrain
and understand the effects of large-scale environmental perturbations on marine settings.
64
CHAPTER IV
Permian-Early Jurassic trends in bryozoan colonial morphology
Abstract
An examination of the relative abundance and paleoenvironmental histories of
bryozoan colonial morphotypes during the Permian, Triassic, and Early Jurassic indicates
a gradual shift in the overall morphological composition of bryozoan assemblages.
Permian and Early Triassic bryozoan communities were dominated by erect species that
were then slowly replaced during the remainder of the Triassic and Early Jurassic by
smaller encrusting species. This morphological switch mirrors a concurrent taxonomic
turnover during which all but one stenolaemate order became extinct by the end of the
Triassic, leaving encrusting members of Order Cyclostomata to dominate the Early
Jurassic. Although bryozoan colonial morphology is often controlled by extrinsic
environmental factors, particularly at the local scale, it is difficult to consider this
morphological turnover independent of the taxonomic turnover, particularly because the
bryozoan taxa that disappeared during this interval (Order Fenestrata, Order
Trepostomata, Order Cryptostomata) were mostly composed of erect colonies. Results
presented in Chapter III showed that long-term environmental degradation associated
with the end-Permian and end-Triassic mass extinctions drove the bryozoan taxonomic
evolution, and must therefore indirectly have caused this morphological change.
65
1. Introduction
Fossil and modern bryozoan colonies are distinguishable from other colonial
invertebrates (i.e., corals, sponges) by their zooids, smaller colony size, and unique
growth forms. Bryozoan colonial morphotypes, which have not changed significantly
since the appearance of Phylum Bryozoa in the Early Ordovician, are not specific to a
particular bryozoan group. Almost every bryozoan genus, regardless of its higher-order
taxonomic affiliation, can display each of the common colony forms. The distribution of
these forms across communities is controlled by environmental and ecological factors
(Hageman et al., 1997 and references within; McKinney and Jackson, 1989). Therefore
changes in the frequency of colonial morphotypes may reflect changes in the
environment.
As discussed in previous chapters, the Permian to Jurassic interval includes three
major extinctions (end-Guadalupian, end-Permian, and end-Triassic mass extinctions)
and two periods of prolonged environmental stress that are clearly reflected in the
paleoenvironmental distribution of bryozoan genera during that interval (see chapters II
& III). In view of that, the frequency and distribution of bryozoan colonial morphotypes
within communities may also reveal the nature of the environmental degradation caused
by the three extinctions.
In this study, I examined the frequency and paleoenvironmental distribution of
bryozoan colonial morphotypes during the Permian, Triassic, and Early Jurassic to
determine 1) the dominant bryozoan colony growth form in each environment for each
time period, if any, and 2) whether extinction-related environmental stress controlled the
66
spatial distribution of each common growth form. These results show whether bryozoan
morphological data, regardless of the taxonomic affiliation of the studied specimens, can
be used to reveal intervals of environmental stress. Given the difficulty of systematically
classifying fossil bryozoans, which leads many researchers to disregard them, the method
tested in this study would make the extraction and application of fossil bryozoan data
more approachable by non-specialists.
2. Previous work
The broad interest from ecologists, sedimentologists, and paleontologists in
understanding the correlation between bryozoan colonial morphology and environmental
conditions stems from the important ecological and sedimentological contribution of
bryozoans to many fossil and recent shelf communities and the rock record (i.e.,
Boardman and Cheetham, 1987; Cuffey, 1977). Bryozoans are ubiquitous in benthic
communities throughout the Phanerozoic and are significant skeletal carbonate
contributors to the sedimentary record (Nelson et al., 1988; Taylor and Allison, 1998; i.e.,
Wass et al., 1970). Several studies have suggested that the composition and diversity of
bryozoan assemblages could provide information such as water depth and substrate
composition (i.e., Kuklinski et al., 2005; Pachut et al., 1995); however the most
promising correlation between habitat conditions and bryozoan diversity also took into
account the relative abundance of bryozoan colonial morphotypes (i.e., Taylor, 2005).
The potential of bryozoan growth forms as environmental indicators was first
reported in a study of Cenozoic cheilostomes that showed a distinct relationship between
67
growth forms and habitat, the latter controlled by water depth and agitation (Stach, 1936).
Subsequent studies using fossil and modern specimens continued to explore this
relationship, often using more detailed growth types in an effort to identify extrinsic
factors that would be applicable across all bryozoan communities and localities (i.e.,
Amini et al., 2004; Bianchi et al., 1991; Cuffey, 1967; Gordon, 1987; Hageman et al.,
1997; Kelley and Horowitz, 1987; Labracherie, 1973; Lagaaij and Gautier, 1965; Nelson
et al., 1988; Pachut and Cuffey, 1999; Pedley, 1976; Taylor, 2005). The overall
conclusions of these papers bolster the relationship between bryozoan morphotypes and
habitat, yet show that: 1) no growth form is confined to single habitats; 2) dominance in
certain habitats is only achieved under optimal conditions; and 3) specific environmental
controls are actually not uniformly applicable (Gordon, 1987; Hageman et al., 1997;
Lagaaij and Gautier, 1965; Taylor, 2005). However, Nelson et al. (1988) demonstrated
that a combination of the association of various growth forms in each habitat and their
relative abundance can yield the most valuable information.
3. Methods
I compiled a global database of the morphology and environmental setting of
Permian, Triassic, and Early Jurassic marine stenolaemate bryozoan species from 108
assemblages (Appendix E). These assemblages represent the sub-sample of assemblages
from Appendix A for which morphological data at the species-level could be extracted.
For this study, morphology refers to the visible, exterior colonial growth form of
bryozoan species. Over the years, many terms stemming from different classification
68
methodologies have been used in the literature to describe bryozoan growth forms. Some
of these classifications are very detailed as they attempt to capture as much ecological
information as possible in a local setting where environmental conditions are already
known (Hageman et al., 1998; i.e.. Lagaaij and Gautier, 1965; McKinney and Jackson,
1989; Smith, 1995). Other classification schemes are linked to the taxonomic assignment
of the studied specimens (i.e., adeoniform is used to describe members of genus
Adeonellopsis) and are difficult to use by non-specialists. (i.e, Brood, 1972; Schopf,
1969; Stach, 1936). Since the goals of this research are to document long-term
morphological trends and to demonstrate the feasibility of this methodology to non-
specialists, I adopted a simple but descriptive classification that can be easily applied and
is not hindered by taxonomic affiliation. Bryozoan species morphology was therefore
categorized as erect, massive, or encrusting (Fig. 1.6). Erect colonies were further
subdivided into ramose (which includes the bifoliate forms) and fenestrate. The collected
data were plotted to examine the relative abundance of each type of growth form through
the Permian to Early Jurassic interval, their relative diversity according to their
taxonomic affiliation at the ordinal level, their environmental distribution, and their
relative abundance in each environment.
4. Relative abundance and taxonomic distribution of bryozoan growth forms
The relative abundance of bryozoan colony morphologies during the Permian to
Jurassic interval indicates that the erect forms dominated across the entire interval,
although not uniformly (Fig. 4.1). Fluctuations in the abundance of erect forms coincide
69
Figure 4.1. Permian to Early Jurassic relative abundance (%) of bryozoan growth forms (ramose + bifoliate, fenestrate,
massive, encrusting).
70
with significant paleoecological events such as the end-Permian and end-Triassic mass
extinctions and the return to normal marine conditions in the aftermath of each. During
the gradual increase in marine environmental degradation in the Late Permian, erect
forms proliferated while all other forms slowly disappeared. Erect bryozoans also
dominated during the Early Triassic recovery but were replaced by more morphologically
simple forms (encrusters and massive forms) once normal marine communities were re-
established in the Middle Triassic. Conversely, erect bryozoans disappeared at the end of
the Triassic and were absent during the recovery period of the Early Jurassic, the latter
being instead characterized by simple encrusters.
Within erect bryozoans, the fenestrate form was the most abundant throughout the
Permian but seemingly vanished from the fossil record of stenolaemate bryozoans at the
end of the Permian. The fenestrate morphology is not observed again in the bryozoan
fossil record until the Cretaceous, first within a group of gymnolaemate bryozoans and
later within the cyclostomes, the sole surviving order of stenolaemates (McKinney, 1986;
Taylor and Larwood, 1988; Walter, 1972).
Taxonomically, the erect morphology was the most common among orders
Trepostomata (Permian: 45%; Triassic: 42%), Cryptostomata (Permian: 61%, Triassic:
100%), and Fenestrata (Permian: 100%) throughout the Permian and Triassic (Fig. 4.2).
Order Cystoporata seemed to be dominated by erect forms during the Permian (34%)
though the lack of data (the colony form of 45% of Permian cystoporates is unknown)
raises doubts about this observation. However, cystoporates were clearly composed of
morphologically simple colonies during the Triassic (Fig. 4.2). Finally, most bryozoans
71
Figure 4.2. Growth form summary for each bryozoan order during the Permian, Triassic, and Jurassic.
72
of Order Cyclostomata were encrusters during the Triassic and Jurassic, and included the
highest percentage of encrusters (33%) from among all other bryozoan orders during the
Permian.
Overall, regardless of taxonomic affiliation, there is a clear progression in the
relative distribution of the major bryozoan growth forms throughout the Permian to
Jurassic interval as erect forms lost their dominance and were gradually replaced by
massive colonies and encrusters (Figure 4.2).
5. Paleoenvironmental distribution of bryozoan growth forms
The environmental distribution of colonial morphotypes through the Permian to
Early Jurassic interval is shown as presence/absence data (Figs 4.3-4.5). Erect forms were
present in all environments during the Permian (Figs. 4.3), although fenestrates account
for most of the diversity in offshore, reef, and mound settings (Fig. 4.6). Erect bryozoans
(absent fenestrates) remained onshore during the Triassic and were restricted to mid-outer
shelf settings during the Jurassic (Fig. 4.3A). Massive colonies and encrusters show no
change in trend throughout the entire interval, largely remaining in onshore settings (Figs.
4.4 and 4.5). A breakdown of the relative abundance of growth forms within each
environment shows that massive and encrusting colonies were completely absent from
the deepest waters during the Permian and Triassic (Fig. 4.6).
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Figure 4.3. Paleoenvironmental history of Permian-Early Jurassic marine stenolaemate
bryozoan erect (A = ramose and bifoliate, B = fenestrate) morphotypes. Each dot
represents a data point, grey boxes indicate the presence of the growth form in an
assemblage, white boxes with dots indicate the absence of the growth form in an
assemblage, and blank boxes indicate the absence of data. E = Early, M = Middle, L =
Late, NS = Nearshore, IS & L = Inner Shelf & Lagoon, MS = Middle Shelf, R = Reefs,
OS = Outer Shelf, SM = Slope Mounds, S & DB = Slope & Deep Basin.
74
75
Figure 4.4. Paleoenvironmental history of Permian-Early Jurassic marine
stenolaemate bryozoan massive morphotypes. See Fig. 4.3 for explanation
of abbreviations.
76
Figure 4.5. Paleoenvironmental history of Permian-Early Jurassic marine
stenolaemate bryozoan encrusting morphotypes. See Fig. 4.3 for
explanation of abbreviations.
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Figure 4.6. Permian to Early Jurassic relative abundance of bryozoan
growth forms for each of the seven environments used in figures 4.3-4.5.
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6. Effects of Late Permian and Late Triassic environmental stress on growth form
trends
The results presented above show a clear long-term trend in the overall
morphological composition of bryozoan communities as they transitioned from the
Permian to the Triassic. McKinney and Jackson (1989) had already observed the
transition from Paleozoic erect-dominated bryozoan communities to communities
dominated by encrusters during the Mesozoic and Cenozoic. However they lacked data
for Late Permian, Triassic, and Early Jurassic species, hindering any detailed analysis
regarding the timing of this transition (Fig. 4.7A). The missing data, provided by this
study reveals that the switch from erect to encrusting species was a gradual process that
initiated during the recovery period of the Early Triassic and culminated with the end-
Triassic extinction (Fig. 4.7B).
McKinney and Jackson (1989) speculated that the proportional decrease of erect
forms in the post-Jurassic was caused by an increase in predation pressure. Cretaceous to
Recent bryozoan colonies show evidence of circular boreholes and anti-predatory
adaptations (spines, superstructures) but Jurassic and older colonies do not (McKinney et
al., 2003; Taylor, 1982; Taylor and Ernst, 2008). Given that the dominant growth form
changed from morphologically complex erect colonies to morphologically simple
encrusters, it is plausible that the change was driven by the long-term environmental
instability characteristic of this time interval, starting with the end-Permian extinction and
lasting through the recovery during the Early Jurassic. Late Triassic and early Jurassic
bryozoan encrusters were small, short-lived ‘weeds’ that may have been more likely to
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Figure 4.7. A = Phanerozoic relative abundance of bryozoan growth forms (erect,
encrusting, and free-living), modified from McKinney and Jackson (1989). Time scale
key: MO = Middle Ordovician, UO = Upper Ordovician, S = Silurian, LD = Lower
Devonian, MD = Middle Devonian, UD = Upper Devonian, LC = Lower Carboniferous,
UC = Upper Carboniferous, LPm = Lower Permian, UPm = Upper Permian, M-UJr =
Middle-Upper Jurassic, LK = Lower Cretaceous, UK = Upper Cretaceous, P =
Paleogene, N-R = Neogene to Recent. B = Late Permian to Early Jurassic relative
abundance of erect and encrusting bryozoan growth forms. There is no record of free-
living bryozoans during that interval.
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survive in environmentally stressed settings (e.g., anoxic, low productivity) (Knoll et al.,
2007; Okamura et al., 2001; Taylor and Ernst, 2008). Carbon isotopic data from several
sections spanning the end-Triassic extinction indicate a productivity collapse associated
with rapid plankton extinction and the presence of regional surface water anoxia
(Ciarapica, 2007; Sephton et al., 2002; Van de Schootbrugge et al., 2007; Ward et al.,
2004; Ward et al., 2001).
However, the patterns observed during each extinction interval are markedly
different; erect species dominated during the End-Permian event while encrusters were
most prominent during and after the end-Triassic extinction, suggesting that the nature of
the environmental stress differed for each extinction. Indeed, differential environmental
trends exhibited by bryozoan genera were also observed at the Permian-Triassic and
Triassic-Jurassic boundaries (see Chapter II), supporting varied causes for these
extinctions. The contrasting morphological compositions of the end-Permian and end-
Triassic extinction could also be related to the taxonomic affiliation of the genera present.
Trepostome bryozoans, which typically grow as erect colonies, dominated the Late
Permian and Early Triassic but died out by the end of the Triassic (Powers and Pachut,
2008; Taylor and Larwood, 1988). The dominant encrusters of the Late Triassic and only
bryozoan group of the Early Jurassic were the cyclostome bryozoans (Powers and Pachut,
2008; Taylor and Ernst, 2008). Environmental conditions undoubtedly played a role in
the distribution of growth form through the Permian to Jurassic interval, either directly by
controlling the growth of individual colonies, or indirectly through the selective survival
of cyclostomes.
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7. Conclusions
The relative abundance of bryozoan colonial morphotypes through the end-
Permian and end-Triassic interval indicates that a gradual large-scale switch between
erect-dominated Paleozoic bryozoan communities and encruster-dominated Mesozoic
communities took place during the Triassic. This transition may have been driven by
extinction-related environmental stress, but may also be a result of the taxonomic
affiliation of the surviving taxa.
This study also shows that bryozoan colony morphological data has the potential
to reveal periods of environmental stress on a global scale through the identification of
stage-level time intervals dominated by a single growth form. However, given the
existing relationship between taxonomic affiliation and growth form, the latter should not
be used exclusively, but instead as a tool to corroborate similar findings attained through
different approaches. This method is easy to use by non-specialists but has several
drawbacks. It may not be applicable on shorter time-scales as only intervals surrounding
major mass extinctions displayed any discernable patterns. For example, the end-
Guadalupian extinction had no effect on the distribution or relative abundance of
bryozoan colony forms. Applying this method on a local, rather than regional or global
scale, may also prove ineffective as individual assemblages vary widely in their
morphological composition and show no clear correlation between environmental setting
and relative abundance of colony forms.
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CHAPTER V
Behavior of lophophorates during the end-Permian mass extinction and recovery
Abstract
The end-Permian mass extinction devastated most marine communities and the
recovery was a protracted event lasting several million years into the Early Triassic.
Environmental and biological processes undoubtedly controlled patterns of recovery for
marine invertebrates in the aftermath of the extinction, but are often difficult to single-
out. The global diversity and distribution of marine lophophorates during the aftermath of
the end-Permian mass extinction indicates that stenolaemate bryozoans, rhynchonelliform
brachiopods, and lingulid brachiopods displayed distinct recovery patterns.
Bryozoans were the most susceptible of the lophophorates, experiencing relatively
high rates of extinction at the end of the Permian, and becoming restricted to the Boreal
region during the Early Triassic. The recovery of bryozoans was also delayed until the
Late Triassic and characterized by very low diversity and abundance. Following the final
disappearance of Permian rhynchonelliform brachiopod survivors, Early Triassic
rhynchonelliform brachiopod abundance remained suppressed despite a successful re-
diversification and a global distribution, suggesting a decoupling between global
taxonomic and ecological processes likely driven by lingering environmental stress.
In contrast with bryozoans and rhynchonelliforms, lingulid brachiopods
rebounded rapidly, colonizing shallow marine settings left vacant by the extinction.
Lingulid dominance, characterized by low diversity but high numerical abundance, was
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short-lived and they were once again displaced back into marginal settings as
environmental stress changed through the marine recovery. The presence in lingulid
brachiopods of the respiratory pigment hemerythrin, known to increase the efficacy of
oxygen storage and transport, when coupled with other morphological and physiological
adaptations, may have given lingulids a survival advantage in environmentally stressed
Early Triassic settings.
1. Introduction
The documentation of global faunal patterns during extinction intervals and their
subsequent survival and recovery phases provides invaluable insight on the processes
driving ecological catastrophes. Although some studies have tried to correlate extinction
patterns with attributes such as geographic range (Jablonski, 1986) or larval types
(Valentine and Jablonski, 1986), most have often focused on the response of individual
groups without incorporating information about their ecological and environmental
context (i.e., Kier, 1973; McGowan, 2004; Pan and Erwin, 2002; Taylor and Larwood,
1988). Assessing the ecological behavior of related groups of marine invertebrates in the
immediate aftermath of mass extinctions offers new opportunities to identify attributes,
whether ecological or biological, that may have helped certain taxa to survive and thrive
during times of increased environmental stress.
By means of recently published data, I reviewed and compared the environmental
distribution and ecological attributes of marine brachiopods and stenolaemate bryozoans
during their recovery from the end-Permian mass extinction. Both groups are sessile
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members of the Paleozoic Fauna (Sepkoski, 1981) and share a lophophorate filter-feeding
life habit, passive respiratory system, and low basal metabolic rate (Knoll et al., 1996).
Along with suspension feeding crinoids and corals, lophophorates were preferentially
affected by the end-Permian extinction event (Knoll et al., 1996; Knoll et al., 2007).
Regardless, several brachiopod and bryozoan taxa survived into the Early Triassic and
ultimately re-diversified. Given their similar physiology and ecological requirements,
Early Triassic brachiopods and bryozoans might be expected to have analogous recovery
patterns. A decoupling of these patterns could indicate that factors controlling the
survival and recovery of marine invertebrates in the wake of the largest Phanerozoic
extinction were more complex than originally inferred.
2. Lophophorates
The lophophorates are a sessile, filter-feeding group of aquatic invertebrate
organisms that includes Bryozoa and Brachiopoda, defined by the use of an extensible
cilia-bearing organ, the lophophore, for feeding and respiration. Members of both
lophophorate phyla are abundant in marine settings today and share a long and rich fossil
record from level-bottom communities; brachiopods first appeared in the Cambrian and
bryozoans appeared in the Ordovician (Taylor and Ernst, 2004; Williams et al., 1996).
Phylum Bryozoa consists of two marine classes, Stenolaemata and
Gymnolaemata, and the freshwater class Phylactolaemata. Stenolaemate and
gymnolaemate bryozoans were important contributors to Phanerozoic marine diversity,
but the diversity of gymnolaemates prior to the Jurassic was negligible. Paleozoic and
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Triassic bryozoan faunas were dominated by five orders, only one of which went extinct
at the end of the Permian (Fig. 5.1) (Taylor and Larwood, 1988). Brachiopods are more
taxonomically diverse and are grouped into the three subphyla Rhynchonelliformea,
Linguliformea, and Craniiformea. Rhynchonelliforms, the largest group with 19 orders,
dominated most post-Ordovician Paleozoic brachiopod faunas and lost three orders
during the end-Permian extinction interval (Fig. 5.1) (Williams et al., 2000).
Bryozoans and brachiopods differ in their morphology. Bryozoans are colonial
organisms composed of genetically identical functional units called ‘zooids’ that secrete a
calcium carbonate skeleton in most species. They have a variety of growth forms
controlled by the arrangement of the zooids and their environmental setting (Hageman et
al., 1998). Conversely, brachiopods are solitary shelled organisms. Unlike bivalves, their
valves are bilaterally symmetric and most live attached to hard substrates on the seafloor
using a fleshy appendage called a pedicle. The Lingulidae (Order Lingulida, subphylum
Linguliformea) are the only infaunal group of brachiopods (Emig, 1997).
3. Early Triassic stenolaemate bryozoans
Stenolaemate bryozoan diversity steadily declined throughout the Late Permian,
culminating in the extinction of 73.5% of stenolaemate genera (Powers and Pachut,
2008), including all members of the Order Fenestrida (Schäfer and Fois, 1987; Taylor
and Larwood, 1988). Early Triassic bryozoan faunas were depauperate, geographically
restricted, and solely composed of trepostomes; taxa belonging to the other surviving
orders did not reappear until the Middle and early Late Triassic (Powers and Pachut,
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Figure 5.1. Schematic illustrating the range of stenolaemate bryozoans
and rhynchonelliform brachiopods through the Permian/Triassic
extinction interval. Data from Powers & Pachut (2008) and Williams et
al. (1996).
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2008). Induan bryozoan faunas were composed of three species belonging to Paleozoic
trepostome survivors Pseudobatostomella and Paralioclema. These species are reported
from nearshore and multiple offshore habitats at three locations in Svalbard and one
location in Siberia (Lazutkina, 1963; Nakrem and Mørk, 1991). Paleogeographic
reconstructions of the Early Triassic indicate that these faunas were restricted to the
Boreal region along the northwestern coast of Pangea (Fig. 5.2). Diversity increased in
the Olenekian with the appearance of two new Triassic genera Arcticopora in the Boreal
region and Dyscritellopsis in eastern Panthalassa (Fritz, 1962; Schäfer et al., 2003). To
date, no Early Triassic bryozoans have been documented from the Tethys Ocean. The
recovery of bryozoans continued into the Middle Triassic when they increased their
geographic range to several localities around the world, including within the Tethys
Ocean. However, bryozoan assemblage generic richness and numerical abundance
remained depleted throughout the Early and Middle Triassic and only gradually increased
during the Late Triassic (Powers and Bottjer, 2007). The recovery of bryozoans was
considered complete with the rapid diversification of genera and species in the early Late
Triassic, coupled with an expansion across shallow onshore to deep offshore
environments (Powers and Bottjer, 2007; Powers and Pachut, 2008).
4. Early Triassic brachiopods
An estimated 87-90% of brachiopod genera became extinct at the end of the
Permian and the surviving rhynchonelliform genera died-out in the early Induan (Carlson,
1991; Chen et al., 2005b; Shen and Shi, 2002). Early Triassic brachiopod assemblages
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Figure 5.2. Paleobiogeographic distribution of Early Triassic bryozoans.
White stars: Induan assemblages (Siberia and Svalbard); Black stars:
Olenekian assemblages: Ellesmere Island, western United States, and
Svalbard. Modified from Powers & Pachut (2008).
90
are composed of three elements – Permian rhynchonelliform brachiopod survivors, new
“Mesozoic-type” brachiopods, and lingulids (Boyer et al., 2004; Chen et al., 2005a, b;
Rodland and Bottjer, 2001).
4.1. Permian survivors
Some of the earliest faunas documented from strata immediately above the
extinction horizon are composed of rhynchonelliform brachiopods that survived the
extinction. They include 102 species belonging to 43 genera dominated by productids,
orthids, spiriferids, and orthotetids (Chen et al., 2005b). These faunas are recorded from
multiple localities in the Tethyan, Peri-Gondwanan, and Panthalassan regions (see Fig. 1
in Chen et al., 2005b), and were most abundant in open platform and offshore settings.
The survival of these Permian rhynchonelliform brachiopods has been attributed to their
small size, cosmopolitan behavior, and broad environmental tolerances (Chen et al.,
2005b). However most died off rapidly during the second extinction pulse associated
with the end-Permian event, less than a million years following the first pulse at the
boundary (Xie et al., 2005).
4.2. Rhynchonelliform brachiopods
The taxonomic recovery of rhynchonelliform brachiopods, defined by the
origination of new higher taxa, was initiated in the mid- to late Induan and continued
throughout the Olenekian (Chen et al., 2005a). The new fauna, composed of 32 species
from 20 genera of rhynchonellids, terebratulids, spririferinids and athyridids, is reported
91
from various locations around the Tethyan, Boreal, and Gondwanan regions (Fig. 5.3)
(Chen et al., 2005a). Despite their wide geographic distribution, Early Triassic
rhynchonelliform brachiopods exhibited a high degree of endemism and several
biogeographic provinces can be distinguished (Chen et al., 2005a). The timing of the
recovery of rhynchonelliform brachiopods between these regions was not simultaneous
and appeared to have happened in two steps. A preliminary recovery took place in the
mid- to late Induan in South China; however these genera died off rapidly and failed to
initiate a successful re-diversification. More successful genera evolved in several other
regions during the late Induan and early Olenekian, spurring the rapid diversification of
more than fifty brachiopod genera during the Middle Triassic (Chen et al., 2005a; Dagys,
1993; Hallam and Wignall, 1997). Early Triassic rhynchonelliform brachiopods were
nonetheless rarely abundant in marine communities, which were instead dominated by
abundant bivalves and gastropods (Clapham and Bottjer, 2007b; Fraiser and Bottjer,
2004, 2007b).
4.3. Lingulid brachiopods
Lingulids, represented by the genus Lingularia, proliferated during the
Griesbachian (Xu and Grant, 1994). They were numerically abundant and dominated
many onshore (nearshore to middle shelf) communities from several regions (South
China, Western Australia, East Greenland, western United States, Japan, Pakistan,
Europe, Iran, Russia) (Fig. 5.4) (Boyer et al., 2004; Chen et al., 2005b; Fraiser and
Bottjer, 2005; Peng et al., 2007 and references therein; Schubert and Bottjer, 1995).
92
Figure 5.3. Early Triassic rhynchonelliform brachiopod faunas. New
Early Triassic brachiopod genera first evolved in South China in the late
Induan (white star) but rapidly went extinct. More successful genera
evolved during the Olenekian at several locations (black stars), initiating
the re-diversification of rhynchonelliforms. Modified from Chen et al.
(2005a).
93
Figure 5.4. Paleobiogeographic distribution of lingulid brachiopod
assemblages during the early Induan. Modified from Peng et al. (2007).
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Early Triassic lingulids were commonly associated with the equally dominant bivalves
Claraia and Promyalina and have been considered both a disaster taxon and an
ecological opportunist (Rodland and Bottjer, 2001; Zonneveld et al., 2007). Disaster taxa,
which normally thrive in stressed marginal environments and are characterized by a long
evolutionary history, frequently colonize and dominate normal marine settings left vacant
in the wake of an environmental crisis before being displaced again as the recovery
progresses (Bottjer, 2001; Hallam and Wignall, 1997). Conversely, ecological
opportunists are regular components of normal marine communities which then
proliferate when other more dominant taxa become extinct during mass extinction
intervals (Harries et al., 1996; Zonneveld et al., 2007). In the case of the end-Permian
mass extinction, lingulids rapidly occupied and proliferated in the newly vacated eco-
space. The prominence of lingulids in Early Triassic communities was short-lived; their
abundance had declined significantly by the end of the Griesbachian as the recovery of
marine organisms progressed and environmental conditions ameliorated (Fraiser and
Bottjer, 2005; Rodland and Bottjer, 2001; Schubert and Bottjer, 1995).
5. Recovery patterns, timing, and tempo
Despite their similar physiology and ecological requirements, rhynchonelliform
brachiopods and stenolaemate bryozoans displayed distinctive patterns of survival and
recovery from the end-Permian mass extinction (Fig. 5.5).
The first rhynchonelliform brachiopod and bryozoan faunas documented in the
immediate aftermath of the end-Permian extinction were composed of survivor genera.
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Figure 5.5. Early Triassic timescale showing the range and characteristics of
rhynchonelliform brachiopod, lingulid, and stenolaemate bryozoan faunas during the
recovery interval of the end-Permian mass extinction. L P: Late Permian; G:
Griesbachian; M Tr: Middle Triassic.
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But whereas the rhynchonelliform brachiopod survivors died shortly after the extinction
interval (~0.7 Ma, Chen et al., 2005b), bryozoan survivor genera Paralioclema and
Pseudobatostomella continued to diversify during the Triassic and did not become extinct
until the Late Triassic. Trepostomes were in fact the most successful bryozoans in the
Triassic, accounting for 81% of all reported Triassic stenolaemate species (Powers and
Pachut, 2008; Schäfer and Fois, 1987).
New rhynchonelliform brachiopod and bryozoan genera appeared during the late
Induan and the Olenekian seeding the recovery and successful re-diversification of both
groups. However, the timing and tempo of their recovery differed. The taxonomic
recovery phase of articulate brachiopods was initiated in the late Induan and completed
by the Middle Triassic. Conversely, bryozoans recovered more gradually during the Early
and Middle Triassic, although they remained uncommon in most Triassic communities.
Ecologically, rhynchonelliform brachiopod abundance rose during the Middle and Late
Triassic, but their breadth across marine communities was more restricted than during the
Paleozoic and continued to decline throughout the remainder of the Phanerozoic
(Clapham et al., 2006). Additionally, rhynchonelliforms became subordinate members of
Mesozoic and Cenozoic communities now dominated by mollusks (see Fig. 2 in Fraiser
and Bottjer, 2007b).
Although the success of several bryozoan Paleozoic holdovers seems to indicate
otherwise, differences in the ecological behavior of bryozoans and brachiopods and a
decoupling in the timing and tempo of their recoveries suggest that bryozoans may not
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have been as well equipped as brachiopods to cope with the environmental effects of the
end-Permian mass extinction.
6. Biological and ecological attributes for survival
In a review of extinction intensity and selectivity among marine invertebrates at
the end of the Permian, Knoll et al. (2007) placed rhynchonelliform brachiopods and
stenolaemate bryozoans in the most vulnerable category, based on their tendency to
secrete robust calcium carbonate skeletons with minimal physiological buffering.
Organisms from this group, that also includes suspension-feeding rugose corals and
crinoids, lost 86% of their genera during the end-Permian extinction, as compared to 5%
for the group with no or little calcium carbonate in their skeleton, to which lingulids
belong. Knoll et al. (2007) argued that elevated concentrations of CO
2
in the photic zone
during the Late Permian would have induced hypercapnic stress in marine invertebrates,
reducing the ability of their respiratory pigments to carry oxygen and disrupting the
biomineralization of their skeletons through oceanic acidification (Caldeira and Wickett,
2003; Fraiser and Bottjer, 2007a; Knoll et al., 1996; Knoll et al., 2007; Raven et al.,
2005). Marine invertebrates like stenolaemate bryozoans and rhynchonelliform
brachiopods that secrete massive skeletons and shells and are unable to buffer against
changes in seawater chemistry would have therefore been unable to survive and thus
experienced high rates of extinction during the end-Permian event. Yet, the difference
between their extinction rates (about 20%) and the duration of their recovery (one vs.
multiple stages) implies that other biological (i.e., pre-adaptations, skeletal mineralogy,
99
morphological complexity, bacterial chemosymbiosis, or reproductive strategies) or
ecological (i.e., environmental range) factors influenced the survival of these
lophophorates during the Permian-Triassic interval (for more in-depth discussion of
attributes of selectivity during extinction see Anstey, 1978; Harries et al., 1996;
Jablonski, 1986; Kitchell et al., 1986).
Nonetheless the recovery of rhynchonelliform brachiopods and stenolaemate
bryozoans contrasts sharply with the ecological behavior exhibited by lingulids during
the Early Triassic. Lingulids dominated a range of localities and habitats in the immediate
wake of the end-Permian crisis, thriving in areas that were subjected to extreme
environmental stress during the extinction. Induan assemblages with abundant lingulids
have been reported from seven localities (western Australia, East Greenland, Pakistan,
Iran, South China, Italy, and eastern Europe) where sedimentological and geochemical
data support the presence of photic zone anoxic to euxinic conditions through the
Permian-Triassic extinction interval (Grice et al., 2005; Kakuwa and Matsumoto, 2006;
Newton et al., 2004; Summons et al., 2006; Wignall et al., 2005; Wignall and Twitchett,
1996). A few rhynchonelliform brachiopods are able to survive in low oxygen conditions,
but lingulids display a much greater tolerance for anoxic settings and may even be able to
survive sulfidic conditions (Bailey et al., 2006; Brunton, 1982; Kammer et al., 1986;
Knoll et al., 2007). Lingulid brachiopods use the non-heme iron protein hemerythrin to
carry and store oxygen (Hammen et al., 1962; Manwell, 1960a). Hemerythrin is
analogous to the respiratory pigment hemoglobin, but is only known to occur in lingulid
brachiopods and sipunculid, priapulid, and annelid worms (Manwell, 1960a, b). Unlike
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other respiratory pigments, hemerythrin in species of lingulids has a low oxygen affinity
and increases the Bohr effect, facilitating oxygen transport and enabling lingulids to
tolerate low oxygen conditions (Manwell, 1960c). This biological attribute likely
provided lingulids with the necessary mechanism to proliferate during the earliest
Triassic. Peng et al. (2007) ascribed the success of lingulids in the aftermath of the end-
Permian event to several pre-adapted morphological traits (small body size and a
phosphatic shell mineralogy) that they were able to modify further (elongation and
flattening of the shells and reduction in shell thickness) to more efficiently survive in the
decreased oxygen levels of the photic zone. The reduction in shell thickness would have
facilitated oxygen exchange directly through the shell, which in turn would have been
more efficiently distributed by the hemerythrin (Manwell, 1960c; Peng et al., 2007). The
mineralogical composition of lingulid shells provided one further advantage by allowing
lingulids to withstand the elevated concentration of oceanic CO
2
(hypercapnia) during the
end-Permian event (Knoll et al., 1996; Knoll et al., 2007). Lingulid shells are phosphatic
and, unlike the calcareous rhynchonelliform brachiopods and stenolaemate bryozoans,
would not have been affected by the undersaturation of calcium carbonate in Late
Permian and Early Triassic seawaters (Fraiser and Bottjer, 2007a). Hypercapnia is
postulated to be one of the principal sources of environmental stress in shallow marine
communities during the end-Permian extinction and a likely contributor to the patterns of
selectivity documented in many marine groups during that interval (Knoll et al., 1996;
Knoll et al., 2007). Although infaunal tiering was reduced during the Early Triassic
(Ausich and Bottjer, 2002; Pruss and Bottjer, 2004), the typical burrowing behavior of
101
lingulids throughout the Phanerozoic may have put them on an adaptive trajectory that,
coupled with their phosphatic shell mineralogy, led to an enhanced ability to use oxygen
and tolerate H
2
S, which may be the key to their success in anoxic to sulfidic settings
since the Cambrian.
7. Conclusions
The diversity, distribution, and abundance of lophophorates during the end-
Permian extinction and subsequent aftermath reveal distinct patterns of survival and
recovery. Stenolaemate bryozoans were the most susceptible of the lophophorates,
experiencing relatively high rates of extinction at the end of the Permian, followed by
very low diversity and abundance during the Early Triassic. Early Triassic bryozoans
were also restricted to the Boreal region where evidence has shown that environmental
conditions after the mass extinction ameliorated more rapidly (Wignall et al., 1998).
Rhynchonelliform brachiopods seemed slightly less sensitive than bryozoans to the
deleterious environmental conditions of the Permian-Triassic interval, and although only
rarely abundant in Early Triassic communities, they were globally distributed. Finally,
lingulids were the least susceptible to the end-Permian event, proliferating in vacated
Griesbachian subtidal habitats.
The success of lingulids, known for their ability to thrive in stressed marginal
environments often characterized by low oxygen conditions, and the lack of
rhynchonelliform brachiopods and stenolaemates bryozoans, suggest that shallow
subtidal settings in the earliest Triassic were not suited for the development of normal
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marine communities and supports geochemical and sedimentological evidence of
sustained environmental degradation during the Early Triassic (i.e., Payne et al., 2004;
Pruss et al., 2006). Additional evidence of sustained environmental stress during the
earliest Triassic comes from size data of microgastropods and Permian survivor
rhynchonelliform brachiopods (Chen et al., 2005b; Fraiser and Bottjer, 2004; Payne,
2005; Twitchett, 2005). Early Triassic microgastropods were opportunistic taxa that, like
lingulids, proliferated in empty subtidal habitats in the aftermath of the end-Permian
extinction. And the survival, albeit short-lived, of Permian rhynchonelliform brachiopods
during the extinction was attributed to their broad environmental adaptations and small
body size, which may have allowed them to live in settings still affected by anoxia and
euxinia. No reduction in zooid and colony size has been noted for Early Triassic
bryozoans, which may explain why, unlike rhynchonelliform brachiopods survivors,
Permian bryozoan holdovers were not globally distributed.
The re-diversification of rhynchonelliform brachiopods during the Early Triassic
was not accompanied with an increase in their abundance in marine communities,
suggesting a decoupling between taxonomic and ecological processes likely driven by
lingering environmental instability. Divergence between global taxonomic diversity and
abundance at times of elevated environmental stress or in the aftermath of mass
extinctions has been noted in the Late Permian and the Early Cenozoic (Danian Stage)
(Clapham and Bottjer, 2007a; McKinney et al., 1998).
The taxonomic and ecological behavior of lophophorates during the end-Permian
mass extinction interval suggest that the environmental effects of this mass extinction
103
were both protracted and more complex than originally inferred and imply that
decoupling between taxonomic and ecological processes is prevalent during extinction
intervals.
104
CHAPTER VI
Conclusions
1. Summary
I examined the diversity and distribution (environmental and geographical) of marine
stenolaemate bryozoans through the end-Permian and end-Triassic mass extinction to:
- Identify the extent and duration of extinction-related environmental stress on
marine environments
- Document the diversity of bryozoans through the Permian to Early Jurassic
interval, calculate new bryozoan extinction rates for the end-Guadalupian, end-
Permian, and end-Triassic extinctions, and constrain the effect of extinction-
related environmental stress on their taxonomic evolution and environmental
distribution
- Examine the effect of mass extinction-related environmental degradation on the
distribution and relative abundance of bryozoan colonial growth forms across
environments
- Identify possible attributes of extinction survival of marine lophophorates
(stenolaemate bryozoans and rhynchonelliform brachiopods) for the end-Permian
mass extinction
Results have led to a better understanding of the ecological impacts of mass extinctions
on the evolutionary trajectory of marine stenolaemate bryozoans and offer a useful tool
105
for identifying protracted intervals of environmental stress. Results and implications for
each aspect of this research are summarized below.
2. Extinction-related environmental stress
1. The paleoenvironmental history of marine stenolaemate bryozoans shows that
extinction-related environmental stress gradually encroached onto marine shelves
prior to the end-Permian and end-Triassic mass extinctions, affecting offshore
communities first and nearshore communities last.
2. The offshore to onshore direction of the environmental stress corroborates
evidence from earlier studies that some deep-water phenomenon was responsible
for the Late Permian and Late Triassic environmental degradation. The end-
Permian and end-Triassic mass extinctions were therefore part of two prolonged
intervals of environmental stress caused by intrinsic, rather than extraterrestrial,
environmental factors. Indeed, recent paleoecological, sedimentological, and
geochemical evidence combined with the observed paleoenvironmental patterns
of Permian to Early Jurassic bryozoans points to a gradual shallowing of
widespread anoxic to euxinic conditions triggered by massive volcanism and
global warming as the likely culprit for both extinctions
3. Bryozoans became restricted to nearshore settings immediately at the Permian-
Triassic boundary and to middle shelf and reefal settings at the Triassic-Jurassic
boundary. These differential environmental trends suggest a potential varied role
106
for stressful atmospheric and terrestrial perturbations on Late Permian and Late
Triassic shallow marine environments.
4. Tracking the onshore-offshore environmental distribution of marine stenolaemate
bryozoans provides a unique approach to identifying prolonged intervals of
environmentally induced stress and its impact on a range of settings.
3. Impact on bryozoan evolution
5. The Permian to Jurassic generic diversity of stenolaemate bryozoans is
characterized by an abrupt extinction in the Late Permian and low diversity levels
throughout the Triassic and Jurassic. Pre-extinction diversity levels of marine
bryozoan genera were not re-established until the mid-Cretaceous, ~150 m.y.
later.
6. Elevated bryozoan extinction rates during the Late Permian and Late Triassic
were coupled with major changes in their habitats. As their diversity decreased,
bryozoans became restricted to onshore settings. Re-colonization of offshore
environments took place in the aftermath of the end-Permian and end-Triassic
extinctions and was coupled with increases in global generic diversity.
7. New extinctions rates were calculated for the end-Guadalupian (35.3%), end-
Permian (73.5%), and end-Triassic (75%) mass extinctions. The taxonomic
effects of the end-Guadalupian extinction were milder than previously described.
The seemingly high extinction rate for marine bryozoan at the end of the Triassic
was the result of very low standing diversity throughout the Triassic.
107
8. The end-Permian mass extinction remained the largest for bryozoans, drastically
reducing global and assemblage generic diversity and triggering a permanent
change in their paleoenvironmental preferences from nearshore to mid-shelf
settings. This permanent habitat change was concurrent with a taxonomic switch
between stenolaemate and gymnolaemate bryozoans, when all but one
stenolaemate order (Cyclostomata) became extinct during the Late Triassic, thus
ending their 285 million year dominance.
9. Elevated extinction intensity in Late Permian nearshore settings may have been
caused by an increase in terrestrial erosion and sediment influx into the ocean
from a global end-Permian plant extinction and/or by a biocalcification crisis
resulting from elevated atmospheric CO
2
.
4. Bryozoan colonial growth forms
10. Documentation of the relative abundance of the stenolaemate bryozoans major
colonial growth forms shows that Permian to Middle Triassic communities were
dominated by erect forms and Late Triassic to Early Jurassic by simple encrusting
forms.
11. These data indicate that a gradual large-scale switch between erect-dominated
Paleozoic bryozoans communities and encruster-dominated Mesozoic
communities took place during the Triassic, mirroring a concurrent taxonomic
turnover during which all but one stenolaemate order became extinct by the end
108
of the Triassic, leaving encrusting members of Order Cyclostomata to dominate
the Early Jurassic.
12. The changing morphological character of bryozoan communities through the
Permian-Jurassic interval may have been driven by extinction-related
environmental stress, but may also be a result of the taxonomic affiliation of the
surviving taxa.
13. Documenting the relative abundance of bryozoan growth forms through time and
across environmental settings may prove useful for revealing periods of
environmental stress on a global scale, but should not be used exclusively given
the variability of growth forms within bryozoan assemblages.
5. Early Triassic lophophorates
14. Among all lophophorates groups, stenolaemate bryozoans were the most
susceptible, experiencing relatively high rates of extinction at the end of the
Permian, followed by very low diversity and abundance during the Early Triassic.
Bryozoans were also geographically restricted during the Early Triassic and
experienced a gradual recovery that was not complete until the early Late
Triassic.
15. Rhynchonelliform brachiopods seemed slightly less sensitive than bryozoans to
the deleterious environmental conditions of the end-Permian extinction, were
globally distributed during the Early Triassic, and recovered more quickly.
109
16. The most successful group of lophophorates in the Early Triassic were the
lingulid brachiopods who rebounded quickly and dominated shallow marine
settings left vacant by the extinction. Lingulids disappeared quickly as
communities began to recover during the late Early Triassic.
17. Lingulid brachiopods are known to thrive in environmentally stressed marginal
settings so their presence in the earliest Triassic communities indicates that
shallow subtidal settings were not suited for the development of normal marine
communities and supports geochemical and sedimentological evidence of
sustained environmental degradation during the Early Triassic.
18. The presence in lingulid brachiopods of the respiratory pigment hemerythrin,
known to increase the efficacy of oxygen storage and transport, when coupled
with other morphological and physiological adaptations, may have given lingulids
a survival advantage in environmentally stressed Early Triassic settings
19. The varied taxonomic and ecological behavior of lophophorates during the Early
Triassic suggests that the environmental effects of this mass extinction were both
protracted and more complex than originally inferred and imply that decoupling
between taxonomic and ecological processes is prevalent during extinction
intervals.
110
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APPENDIX A
Permian to early Jurassic formations
All Permian, Triassic, and Early Jurassic stratigraphic horizons containing marine stenolaemate bryozoan assemblages
compiled from the literature, taxonomic monographs, and the Paleobiology and PaleoReef Databases (see Appendix B
for a full list of references). Also included are assemblages with rhynchonelliform brachiopods that were used as the
taphonomic control group (indicated by ‘Brachiopoda’ under generic diversity). Age is given at the stage level. From
oldest to youngest, Permian stages include the Asselian, Sakmarian, Artinskian, Kungurian, Roadian, Wordian,
Capitanian, Wuchiapingian, and Changhsingian. Triassic stages include the Induan, Olenekian, Anisian, Ladinian,
Carnian, Norian, and Rhaetian. Early Jurassic stages include the Hettangian, Sinemurian, Pliensbachian, and Toarcian.
Lithostratigraphic unit Location Age Environment
Generic
diversity
‘Oberrhätkalk’ Fm (Dachstein Ls) Austria Norian-Rhaetian Reefs & bioherms Bryozoa indet.
“Unnamed” B Fm Sverdrup basin, Canada Artinskian-Early
Kungurian
Slope and deep basin Bryozoa indet.
Abadeh Fm - lower Unit 4 Iran Capitanian Inner shelf & lagoon 3
Abadeh Fm - upper Unit 4 Iran Capitanian Middle shelf 1
Abdullino Reef Urals, Russia Sakmarian Reefs & bioherms 1
Adnet, Rötelwand, Feichtenstein ,
Gruber Reefs
Austria Rhaetian Reefs & bioherms Bryozoa indet.
Adz’va Fm Urals, Russia Kungurian 2
Aflenz Limestone Austria Norian Reefs & bioherms Bryozoa indet.
Aiduna Fm - Location A Indonesia Early Artinskian Nearshore 14
Aiduna Fm - Location B Indonesia Early Artinskian Nearshore 7
127
Lithostratigraphic unit Location Age Environment
Generic
diversity
Akiyoshi Limestone Gp - Lepidolina
multiseptata shiraiensis zone
Japan Capitanian Inner shelf & lagoon 13
Akiyoshi Limestone Gp - Parafusulina
kaerimizensis zone
Japan Late Kungurian Nearshore-Inner shelf 11
Akiyoshi Limestone Gp - Pseudofusulina
ambigua zone
Japan Sakmarian-
Artinskian
Reefs & bioherms 4
Akiyoshi Limestone Gp - Pseudofusulina
vulgaris zone
Japan Late Asselian Nearshore-Inner shelf 2
Akiyoshi Limestone Gp - Triticites
simplex zone
Japan Asselian Nearshore-Inner shelf 1
Alibashi Fm Iran Changsingian 1
Amaltheen Formation Germany Pliensbachian Middle shelf-Outer shelf 3
Amb Fm - Jan Sukh Mbr Salt Range, Pakistan Wordian Inner shelf & lagoon Bryozoa indet.
Amphyclina Beds Slovenia Carnian Reefs & bioherms 1
Angjie Fm Tibet Autonomous
Region, China
Artinskian Reefs? 13
Antiinskaya Fm Transbaikal region,
Russia
Roadian-Early
Wordian
7
Antimonio Fm Mexico Norian Nearshore-Inner shelf Brachiopoda
Arthurton Gp New Zealand Wuchiapingian Bryozoa indet.
Bai’d Fm Oman Wordian-
Capitanian
Reefs & bioherms Bryozoa indet.
Baker Fm Carnarvon Basin,
Australia
Wordian Outer shelf 4
Balikelik (Baliqliq) Fm Xinjiang Autonomous
Region, China
Late Kungurian-
Roadian
Inner shelf & lagoon 14
Barabash Fm (Barabashevka Fm) Primor’re region, Russia Capitanian 7
Basal Beds Tasmania, Australia Sakmarian 4
Baten beni Zid Series (Lower Biohermal
Complex/Dolomies Inferieures)
Tunisia Capitanian Inner shelf & lagoon 1
Beattie Limestone Fm - Cottonwood Ls
Mbr
Kansas, United States Asselian Bryozoa indet.
Beattie Limestone Fm - Florena Shale
Mbr
Kansas, United States Asselian Bryozoa indet.
Belcher Channel Fm Ellesmere Island, Canada Asselian-
Sakmarian
Inner shelf-Middle shelf 1
128
Lithostratigraphic unit Location Age Environment
Generic
diversity
Belebeevskaia Fm - Petshishiskaia Mbr Urals, Russia Late Wordian Inner shelf & lagoon 3
Belebeevskaia Fm - Prekazan Mbr Urals, Russia Late Wordian Inner shelf & lagoon 2
Belebeevskaia Fm -
Verkhneouslonskaia Mbr
Urals, Russia Late Wordian Inner shelf & lagoon 4
Bellow Fm - Middle Sandstone Mbr Canada Artinskian-
Kungurian
Nearshore Bryozoa indet.
Belloy Fm - Upper Carbonate Mbr Canada Wordian Inner shelf & lagoon? Bryozoa indet.
Berriedale Limestone Tasmania, Australia Artinskian Middle shelf 6
Bioclastic Limestone Oman Induan Outer shelf Brachiopoda
Bird Spring Fm - Unit I Nevada, United States Sakmarian Bryozoa indet.
Bleskovy pramen Fm Slovakia Early Rhaetian Brachiopoda
Blind Fjord Fm Ellesmere Island, Canada Olenekian Slope and deep basin 1
Blue Lias Fm - Porthkerry Mbr England Sinemurian Middle shelf 1
Bohunice Fm Slovakia Kimmeridgian Brachiopoda
Buckskin Mountain Fm Nevada, United States Artinskian-
Kungurian
9
Buffel Fm Bowen Basin, Australia Artinskian Middle shelf 17
Bulgadoo Shale Carnarvon Basin,
Australia
Late Artinskian Slope and deep basin 1
Bundella Mudstone Tasmania, Australia Sakmarian 5
Cadeby Fm - Wetherby Mbr England Wuchiapingian Reefs & bioherms 4
Calcari Grigi Fm Italy Sinemurian Inner shelf & lagoon Bryozoa indet.
Callytharra Fm Carnarvon Basin,
Australia
Sakmarian Inner shelf & lagoon 17
Caodigou Fm Qinghai Province, China Wordian-
Capitanian?
25
Capitan Fm New Mexico, United
States
Capitanian Inner shelf-Reefs-Slope 3
Cassian Fm Italy Early Carnian Slope and deep basin Brachiopoda
Cathedral Mountain Fm Texas, United States Kungurian Slope mounds 12
Cathedral Mountain Fm - Poplar Tank
Mbr
Texas, United States Artinskian Slope mounds 14
Cathedral Mountain Fm - Sullivan Peak
Mbr
Texas, United States Kungurian Slope mounds 6
Cathedral Mountain Fm - Wedin Congl.
Mbr
Texas, United States Kungurian Slope mounds 8
129
Lithostratigraphic unit Location Age Environment
Generic
diversity
Cattle Creek Fm Bowen Basin, Australia Artinskian Inner shelf-Outer shelf 5
Cattle Creek Fm - Eurydesma
Limestone
Bowen Basin, Australia Artinskian 9
Central Urals Urals, Russia Asselian-
Sakmarian-
Artinskian
Middle shelf-Outer shelf Bryozoa indet.
Chandalaz Fm - Okrainsk Terrane Primor’re region, Russia Capitanian Nearshore 14
Chandalaz Fm - Okrainsk Terrane Primor’re region, Russia Capitanian Reefs & bioherms 15
Chandalaz Fm - Voznesensk Terrane Primor’re region, Russia Late Wordian-
Capitanian
Inner shelf-Middle shelf 12
Changhsing Fm - Jiaozishan Section
Bed 25
Guizhou Province, China Changhsingian Middle shelf 7
Changhsing Fm - Member 2 Hunan Province, China Changhsingian Reefs & bioherms 2
Changsing Fm China Changhsingian Onshore Brachiopoda
Charmouth Mudstone Fm England Early Pliensbachian Inner shelf & lagoon? 1
Charmouth Mudstone Fm - Bed Z England Early Pliensbachian Middle shelf 2
Chhidru Fm Salt Range, Pakistan Changhsingian Nearshore 13
Chihsia Fm (Qixia Fm) Sichuan and Guizhou
provinces, China
Kungurian Middle shelf 28
Chumik Fm Himalayas, India Late Sakmarian Bryozoa indet.
Chuping Limestone - Bukit Mata Ayer Malaysia Late Artinskian? Inner shelf & lagoon? 1
Chuping Limestone - Pulau Jonk Malaysia Late Artinskian-
Kungurian
Nearshore 3
Coolkilya Sandstone Carnarvon Basin,
Australia
Roadian Nearshore 1
Copacabana Fm Bolivia Asselian-
Sakmarian
Nearshore 12
Copacabana Fm - Section 2 Cusco Province, Peru Asselian-
Sakmarian
2
Copacabana Fm - Section 7 Apurimac Province, Peru Asselian-
Sakmarian
Middle shelf 1
Copacabana Fm - Section 8 Apurimac Province, Peru Asselian-
Sakmarian
Middle shelf 5
Copacabana Fm - Section 9 Ayacucho Province, Peru Asselian-
Sakmarian
Middle shelf 6
Copper Shale (Kupferschiefer) Fm Poland Changhsingian Inner shelf & lagoon Bryozoa indet.
130
Lithostratigraphic unit Location Age Environment
Generic
diversity
Counsel Creek Fm Tasmania, Australia Artinskian 3
Coyote Butte Fm Oregon, United States Artinskian Bryozoa indet.
Cundlego Fm Carnarvon Basin,
Australia
Late Artinskian Inner shelf-Outer shelf 1
Dalong Fm Zhejiang Province, China Changhsingian Slope and deep basin Brachiopoda
Darlington Limestone Tasmania, Australia Sakmarian Middle shelf-Outer shelf? 1
Deep Bay Fm Tasmania, Australia Late Artinskian 2
Degerböls Fm Ellesmere Island, Canada Wordian-
Capitanian
Inner shelf & lagoon Bryozoa indet.
Dinwoody Fm Montana, United States Induan Middle shelf Brachiopoda
Divya Fm Urals, Russia Upper Artinskian Outer shelf Bryozoa indet.
Dixie Valley Fm Nevada, United States Late Anisian Nearshore 1
Dont Fm Italy Late Anisian Reefs & bioherms 1
Dwyka Fm Namibia Sakmarian Nearshore 1
Dyrham Fm England Pliensbachian Inner shelf-Middle shelf 2
Dzhiakun’skaya Fm Khabarovsk Territory,
Russia
Kungurian 4
Elephant Canyon Fm Utah, United States Asselian Inner shelf-Middle shelf 3
Episkopi Fm Greece Wuchiapingian-
Changhsingian
Inner shelf-Middle shelf Bryozoa indet.
Episkopi Fm Greece Changhsingian Outer shelf 1
Episkopi Fm Greece Changhsingian Reefs & bioherms 1
Fangshankou Fm Gansu Province, China Changhsingian Bryozoa indet.
Fatra Fm Slovakia Rhaetian Brachiopoda
Favret Fm Nevada, United States Anisian Middle shelf 1
Feixianguan Fm Sichuan Province, China Induan Inner shelf & lagoon Brachiopoda
Flowers Fm New Zealand Roadian-Wordian Inner shelf & lagoon? 3
Ford Fm England Wuchiapingian Reefs & bioherms 6
Fossil Cliff Fm Perth Basin, Australia Sakmarian Bryozoa indet.
Fusulinid Limestone Fm Serbia Asselian Slope mounds Bryozoa indet.
Gaptank Fm Texas, United States Asselian Reefs & bioherms Bryozoa indet.
Gerence Fm Turkey Anisian Reefs & bioherms Bryozoa indet.
Gerennavàr Fm - boundary bed Hungary Late
Changhsingian
Outer shelf Brachiopoda
Gerster Limestone Nevada, United States Capitanian Middle shelf-Outer shelf 25
Ghalilah Fm - Lower Mbr United Arab Emirates Norian Inner shelf & lagoon Brachiopoda
131
Lithostratigraphic unit Location Age Environment
Generic
diversity
Gipshuken Fm Spitzbergen, Norway Artinskian Inner shelf & lagoon 2
Great Bear Cape Fm Ellesmere Island, Canada Artinskian Middle shelf-Outer shelf 22
Grenola Limestone Fm - Burr Limestone
Mbr
Kansas, United States Asselian Bryozoa indet.
Grenola Limestone Fm - Legion Shale
Mbr
Kansas, United States Asselian Bryozoa indet.
Grenola Limestone Fm - Neva
Limestone Mbr
Kansas, United States Asselian Inner shelf-Middle shelf Bryozoa indet.
Grenola Limestone Fm - Salem Point
Shale Mbr
Kansas, United States Asselian Bryozoa indet.
Gua Musang Fm Malaysia Early-Middle
Changhsingian
Bryozoa indet.
Gufeng Fm (Kuhfeng Fm) Anhui Province, China Roadian Outer shelf Brachiopoda
Gundara Fm Tajikistan Roadian Reefs & bioherms 1
Gungri Fm Himalayas, India Wuchiapingian Middle shelf Brachiopoda
Güvercinlik Fm Turkey Norian Reefs & bioherms Bryozoa indet.
Hajier Fm Qinghai Province, China Wuchiapingian Inner shelf & lagoon 18
Hambast Fm - Unit 6 Iran Wuchiapingian Outer shelf 2
Hambast Fm - Unit 7 Iran Changsingian Outer shelf Brachiopoda
Hambergfjellet Fm Bjørnøya, Norway Artinskian-Early
Kungurian
Outer shelf 5
Hanwang Fm - Lower Mbr Sichuan Province, China Carnian Nearshore 1
Hanwang Fm - Upper Mbr Sichuan Province, China Carnian Nearshore-Inner shelf-Reefs 1
Hardman Fm - Cherrabun Mbr Canning Basin, Australia Wuchiapingian Nearshore 2
Harsuhai Gp Inner Mongolia, China Changhsingian Onshore Bryozoa indet.
Heshan Fm Guangxi Autonomous
Region, China
Wuchiapingian Outer shelf 2
Hess Fm - Taylor Ranch Mbr Texas, United States Artinskian Middle shelf-Outer shelf Bryozoa indet.
Hetvehely Dolomite Hungary Early Anisian Inner shelf & lagoon Brachiopoda
Hodul Unit - Kasal Limestone Mbr Turkey Norian-Rhaetian Reefs & bioherms Bryozoa indet.
Huai Thak Fm Thailand Changhsingian 1
Hybe Beds Slovakia Rhaetian Nearshore Brachiopoda
Hybe Fm Slovakia Rhaetian Middle shelf 2
Ichinotani Fm Japan Asselian Reefs & bioherms Bryozoa indet.
Iwaizaki Fm - Unit 4 Japan Roadian Nearshore 8
Iwaizaki Fm - Unit 6 Japan Wordian Reefs & bioherms 5
132
Lithostratigraphic unit Location Age Environment
Generic
diversity
Iwaizaki Fm - Unit 8 Japan Capitanian Inner shelf & lagoon 12
Jamal Fm - Bagh-e Vang Mbr Iran Kungurian Reefs & bioherms 5
Juripu Fm Tibet Autonomous
Region, China
Capitanian-
Wuchiapingian
Bryozoa indet.
Kaftarmol Fm Tajikistan Wuchiapingian Reefs & bioherms 1
Kaibab Fm - Alpha Mbr Arizona, United States Roadian Inner shelf & lagoon 3
Kaibab Fm - Beta Mbr Arizona, United States Roadian Inner shelf & lagoon 17
Kalundar Fm Xinjiang Autonomous
Region, China
Wordian-
Capitanian
Inner shelf & lagoon Bryozoa indet.
Kamenetskaia Fm Belarus Wordian 1
Kangshare Fm Tibet Autonomous
Region, China
right below
Permian-Triassic
boundary
Offshore Bryozoa indet.
Kanokura Fm Japan Wordian Inner shelf & lagoon 11
Kapp Dunér Fm Bjørnøya, Norway Asselian Inner shelf & lagoon 1
Kapp Dunér Fm Bjørnøya, Norway Asselian Reefs & bioherms 1
Kapp Starostin Fm - Svenskegga &
Hovtinden Mbrs
Spitzbergen, Norway Roadian-Wordian Middle shelf 15
Kapp Starostin Fm - Svenskegga &
Hovtinden Mbrs
Spitzbergen, Norway Roadian Outer shelf 11
Kapp Starostin Fm - Vøringen Mbr 5y Spitzbergen, Norway Late Artinskian-
Kungurian
Nearshore 22
Karazin Fm Primor’re region, Russia Anisian Outer shelf 1
Kendlbach Fm - Breitenberg Mbr Austria Early Hettangian Inner shelf & lagoon Brachiopoda
Kendlbach Fm - Tiefengraben Mbr Austria Early Hettangian Inner shelf & lagoon Brachiopoda
Khao Pun Fm - Khao Pun Nua Mbr Thailand Wordian Nearshore Bryozoa indet.
Khao Sung Fm - Khao Sung Klang Mbr Thailand Leonardian Nearshore Bryozoa indet.
Khuff Fm Oman Wordian Inner shelf-Outer shelf Bryozoa indet.
Khuff Fm - Unit C Oman Wordian Nearshore Brachiopoda
Khunamuh Fm - Unit E1 Himalayas, India Late
Changhsingian
Deepening Brachiopoda
Kiama Sandstone Mbr Sydney Basin, Australia Wordian Nearshore 1
Kim Fjelde Fm Greenland, Denmark Late Artinskian-
Kungurian
Outer shelf 1
Kistefjellet Fm Spitzbergen, Norway Induan-Olenekian Slope and deep basin 1
Kockatea Shale - Hovea Mbr Perth Basin, Australia Wuchiapingian Inner shelf-Middle shelf Bryozoa indet.
133
Lithostratigraphic unit Location Age Environment
Generic
diversity
Kodiang Limestone Malaysia age too uncertain Redeposition 2
Koguchi Fm Japan Norian Reefs & bioherms Bryozoa indet.
Kössen Facies Czech Republic Norian-Rhaerian Brachiopoda
Kozhim Carbonate Bank Urals, Russia Asselian-
Sakmarian
Reefs & bioherms Bryozoa indet.
Kozhim Fm Urals, Russia Upper Kungurian Middle shelf 3
Kozhim Rudnik Fm Urals, Russia Early Roadian Inner shelf & lagoon 1
Kuling Fm Himalayas, India Wuchiapingian Nearshore Bryozoa indet.
Kuma Fm Japan Capitanian 4
Kunlun Reef Belt Qinghai Province, China Wordian-
Capitanian
Reefs & bioherms Bryozoa indet.
Laborcita Fm New Mexico, United
States
Asselian Inner shelf & lagoon Bryozoa indet.
Laibuxi Fm (Tulung Gp) Tibet Autonomous
Region, China
Ladinian Inner shelf & lagoon 3
Lazurnaya Fm - Beds 4 & 6 Primor’re region, Russia Induan Nearshore Brachiopoda
Lazurnaya Fm - Beds 8 Primor’re region, Russia Early Olenekian Nearshore Brachiopoda
Lehusis Fm Greece Asselian Reefs & bioherms Bryozoa indet.
Lemegou Fm Qinghai Province, China Kungurian-
Roadian?
4
Lengwu Fm Zhejiang Province, China Capitanian Reefs & bioherms 13
Lenox Hills Fm Texas, United States Sakmarian?-
Artinskian
Inner shelf & lagoon Bryozoa indet.
Liangshan Fm Sichuan Province, China Asselian-
Sakmarian
Nearshore 5
Liard Fm Canada Ladinian Reefs & bioherms Bryozoa indet.
Lightjack Fm Canning Basin, Australia Roadian Inner shelf & lagoon 9
Limestone Mbr - Jenka Pass Malaysia Wordian 2
Limestone Mbr - Kampong Awah Quarry Malaysia Wordian 4
Lindstrom Fm Ellesmere Island, Canada Wuchiapingian Inner shelf & lagoon Bryozoa indet.
Loray Fm Nevada, United States Kungurian Middle shelf 2
Lower Bijni Tectonic Unit (Boulder-Slate
Sequence)
Himalayas, India Asselian-? 3
Lower Colina Limestone Mexico Asselian-
Sakmarian
Middle shelf Bryozoa indet.
Lower Fm Thailand Asselian Bryozoa indet.
134
Lithostratigraphic unit Location Age Environment
Generic
diversity
Lower Gechang Fm Himalayas, India Late Sakmarian 1
Lower Lugin Gol Fm Mongolia Roadian Nearshore 2
Lungtan Fm (Luntan/Loping Fm) - 5th
Mbr
Sichuan Province, China Wuchiapingian Bryozoa indet.
Lungtan Fm (Luntan/Loping Fm) -
Xiapeng Mbr
Sichuan Province, China Wuchiapingian Middle shelf 9
Lungtan Fm (Luntan/Loping Fm) -
Xiapeng Mbr
Jiangxi Province, China Wuchiapingian Middle shelf 5
Lungtan Fm (Luntan/Loping Fm) -
Xiapeng Mbr
Guizhou Province, China Wuchiapingian Middle shelf 9
Luning Fm Nevada, United States Norian Inner-Middle-Outer shelf Brachiopoda
Lyons Gp Carnarvon Basin,
Australia
Asselian-Early
Sakmarian
2
Lyudyanza Fm Khabarovsk Territory,
Russia
Wuchiapingian 2
Lyudyanza Fm - Nakhodka Reef Primor’re region, Russia Wuchiapingian-
Early Changsingian
Reefs & bioherms 5
Malbina Fm Tasmania, Australia Early Wordian Outer shelf 2
Mara Fm New Caledonia, Overseas
territory of France
Wuchiapingian Brachiopoda
Marlstone Rock Fm England Toarcian Outer shelf 2
Marlstone Rock Fm England Pliensbachian Outer shelf 2
Marmari Fm Greece Capitanian Nearshore?-Middle shelf? Bryozoa indet.
Merbah el Oussif Series - Djebel Tebaga
Biohermal Complex
Tunisia Capitanian Reefs & bioherms 3
Middle Fm Japan Capitanian Redeposition 7
Miharano Fm Japan Asselian Reefs & bioherms 5
Mikin Fm - Himalayan Muschelkalk Mbr Himalayas, India Anisian-Middle
Ladinian
Middle shelf Brachiopoda
Minnie Point Fm Tasmania, Australia Kungurian-Roadian 1
Miras Fm Greece Changhsingian Inner shelf & lagoon Bryozoa indet.
Miseryfjellet Fm Bjørnøya, Norway Kungurian Nearshore 19
Mission Argillite Fm Washington, United
States
Late Wordian Reefs & bioherms 26
Moenkopi Fm - Virgin Limestone Mbr Utah, United States Olenekian Inner shelf-Middle shelf Brachiopoda
Moribu Fm - Lower Mbr Japan Wordian Bryozoa indet.
135
Lithostratigraphic unit Location Age Environment
Generic
diversity
Morono Fm - Taishaku Limestone Japan Sakmarian 3
Mt Bailey Fm Ellesmere Island, Canada Asselian Slope mounds Bryozoa indet.
Mukut Fm Nepal Anisian-Carnian Outer shelf Brachiopoda
Murdock Mountain Fm Nevada, United States Wordian Outer shelf 10
Murihiku Terrane New Zealand Norian Nearshore 2
Nabeyama Fm (Kuzuu Limestone) -
Yamasuge Limestone Mbr
Japan Artinskian Outer shelf? 4
Nagyvisnyó Limestone Fm Hungary Wuchiapingian-
Changhsingian
Middle shelf Bryozoa indet.
Naizawa Fm Japan Early Carnian Middle shelf 4
Nalbia Sandstone Carnarvon Basin,
Australia
Roadian Inner shelf-Middle shelf 1
Nansen Fm Ellesmere Island, Canada Sakmarian Outer shelf Bryozoa indet.
Naujoji Akmené Fm Lithuania Wuchiapingian 5
Nayband Fm Iran Rhaetian Reefs & bioherms 4
Nelynya-shor Fm Urals, Russia Sakmarian Outer shelf? 3
Nenets Horizon Timan-Pechora basin,
Russia
Asselian 1
Neoschwagerina Beds Slovenia Wordian Reefs & bioherms 9
Nesen Fm Iran Wuchiapingian Inner shelf & lagoon? Brachiopoda
Nimaloksa Fm Himalayas, India Late Carnian-Early
Norian
Nearshore Bryozoa indet.
Nooncanbah Fm Canning Basin, Australia Kungurian Outer shelf 33
Nordenskiölbreen Fm - Tyrrellfjellet Mbr Spitzbergen, Norway Asselian-Early
Sakmarian
Nearshore-Inner shelf 17
Nordenskiölbreen Fm - Tyrrellfjellet Mbr Spitzbergen, Norway Asselian Reefs & bioherms Bryozoa indet.
Novy Svet Fm Slovakia Hettangian Middle shelf Brachiopoda
Nukada Fm - Takauchi Limestone Japan Capitanian 11
Oberrhaet Fm Austria Rhaetian Brachiopoda
Ochiai Fm - Toyazawa Mbr Japan Wordian Bryozoa indet.
Omi Limestone Japan Asselian 5
Omolon Fm Siberia, Russia Roadian-Wordian 6
Oskhtinskaya Fm (Osakhta Fm) Khabarovsk Territory,
Russia
Capitanian Inner shelf & lagoon? 13
Oudjah el Rhar Series ("Calcaires a
Bellerophon")
Tunisia Capitanian Inner shelf & lagoon 3
136
Lithostratigraphic unit Location Age Environment
Generic
diversity
Oxtrack Fm Bowen Basin, Australia Kungurian Inner shelf & lagoon 7
Oxytoma-Mytilus beds Japan Early Carnian Nearshore 1
Pahalgam Fm Himalayas, India Wuchiapingian Middle shelf?-Outer shelf? Bryozoa indet.
Palaeoaplysina mound - Yukon Canada Sakmarian Reefs & bioherms Bryozoa indet.
Palmarito Fm Venezuela Roadian Inner shelf & lagoon 1
Palmarito Fm Venezuela Late Artinskinan Outer shelf 1
Panther Seep Fm New Mexico, United
States
Asselian Middle shelf 4
Pantokrator Fm Greece Carnian Reefs & bioherms 1
Patlanoaya Fm Mexico Asselian Inner shelf & lagoon Bryozoa indet.
Peca-Mezika Reef Slovenia Ladinian Reefs & bioherms Bryozoa indet.
Pha Huat Fm Thailand unknown stage -
Middle Permian
n/a
Pietra di Salomone Megablock Italy Wordian Slope mounds 2
Poduan Fm Guizhou Province, China Anisian Reefs & bioherms 2
Poole Sandstone - Nura Nura Mbr Canning Basin, Australia Early Artinskian Nearshore 9
Puchenpra Fm Nepal Wordian Nearshore 2
Puchenpra Fm - Member C Nepal Capitanian Redeposition 1
Qingyan Fm - Leidapo Mbr Guizhou Province, China Late Anisian Slope and deep basin Bryozoa indet.
Qubuerga Fm Tibet Autonomous
Region, China
Wuchiapingian Nearshore 7
Raanes Fm (Assistance Fm) Ellesmere Island, Canada Artinskian Slope mounds Bryozoa indet.
Raibl Fm - Leckkogel Beds Austria Carnian Reefs & bioherms 1
Rama Fm Himalayas, India Middle Carnian Nearshore Bryozoa indet.
Ranger Canyon Fm Canada Roadian-Wordian Inner shelf & lagoon Bryozoa indet.
Rat Buri Limestone - Khao Chong
Krachok
Thailand Late Sakmarian-
Early Artinskian
4
Rat Buri Limestone – Khao Mang Lat Thailand Late Artinskian Nearshore Bryozoa indet.
Rat Buri Limestone - Khao Phrik Thailand Late Artinskian Outer shelf? 8
Rat Buri Limestone - Khao Raen Thailand Late Artinskian 11
Rat Buri Limestone - Khao Ta Mong Rai Thailand Late Sakmarian-
Early Artinskian
7
Rat Buri Limestone - Ko Muk Island Thailand Late Artinskian Outer shelf 13
Rat Buri Limestone – Phangnga Thailand Artinskian Outer shelf Bryozoa indet.
Recoaro Fm Italy Late Anisian Reefs & bioherms 2
137
Lithostratigraphic unit Location Age Environment
Generic
diversity
Red Eagle Limestone Fm - Bennett
Shale Mbr
Kansas, United States Asselian Bryozoa indet.
Redcar Mudstone Fm England Hettengian Middle shelf 1
Riga Fm Greece Artinskian Nearshore Bryozoa indet.
Road Canyon Fm Texas, United States Late Kungurian-
Early Roadian
Reefs & bioherms 19
Robledo Mountains Fm New Mexico, United
States
Sakmarian Inner shelf & lagoon 6
Rong Kwang Fm Thailand Changhsingian Reefs & bioherms 1
Ruteh Fm Iran Wordian Inner shelf-Middle shelf 4
Safetdaron Fm Tajikistan Kungurian Reefs & bioherms Bryozoa indet.
Saikra Biohermal Complex Tunisia Capitanian Reefs & bioherms Bryozoa indet.
Saiwan Fm Oman Late Sakmarian Nearshore-Inner shelf Bryozoa indet.
San Cassiano Fm Italy Early Carnian Slope mounds 5
Sandy Shale Fm - Mbr 1 Urals, Russia Late Wordian Bryozoa indet.
Sarga Fm Urals, Russia Upper Artinskian Reefs & bioherms 1
Saser Brangsa Fm Himalayas, India Wuchiapingian Slope and deep basin Brachiopoda
Scherrer Fm Mexico Kungurian Inner shelf & lagoon Bryozoa indet.
Schlern Fm Italy Late Anisian Reefs & bioherms 1
Schlern Fm Italy Late Anisian Slope and deep basin 1
Schmidt Fm Primor’re region, Russia Middle Olenekian Nearshore? Bryozoa indet.
Schnoll Fm - Guggen Mbr Austria Middle-Late
Hettangian
Slope and deep basin Brachiopoda
Schnoll Fm - Langmoos Mbr Austria Early Hettangian Slope and deep basin Brachiopoda
Selong Fm – Xishan Section – Coral
Bed
Tibet Autonomous
Region, China
Changhsingian Nearshore Bryozoa indet.
Shaiwa Gp Guizhou Province, China Latest
Changsingian
Slope and deep basin Brachiopoda
Shaktau mound complex (Davansk
Reefs)
Urals, Russia Asselian-
Sakmarian
Reefs & bioherms-Slope
mounds
Bryozoa indet.
Shiyuan Fm Sichuan Province, China Norian Inner shelf & lagoon Brachiopoda
Shoushangou Fm Jilin Province, China Kungurian 7
Skinner Ranch Fm Texas, United States Artinskian-
Kungurian
Inner shelf-Reefs Bryozoa indet.
Skipping Ridge Fm Tasmania, Australia Late Sakmarian 1
Skipping Ridge Fm Tasmania, Australia Early Artinskian 5
138
Lithostratigraphic unit Location Age Environment
Generic
diversity
Sokyuiskaya Fm Transbaikal region,
Russia
Middle Wordian Nearshore 6
Songgui Fm Yunnan Province, China Norian Inner shelf & lagoon 1
Sotong Limestone Malaysia Norian Outer shelf - Slope mounds? Bryozoa indet.
Speiser Shale Kansas, United States Sakmarian Middle shelf 9
Stephens Fm New Zealand Changsingian Brachiopoda
Sterlibashevskala Fm - Baitouganskaia
Mbr
Urals, Russia Early Wordian 1
Sterlibashevskala Fm - Kamishlinskaia
Mbr
Urals, Russia Early Wordian 1
Sunflower Fm - Limestone Mbr Nevada, United States Asselian Slope and deep basin 7
Surmaq Fm - Unit 3 Iran Capitanian Inner shelf-Middle shelf 4
Tak Fa Fm - Khao Hin Kling Thailand Wuchiapingian 5
Tak Fa Fm - Location A Thailand ? 2
Tak Fa Fm - Location K Thailand ? 10
Tak Fa Fm - Location L Thailand ? 1
Tak Fa Fm - Location N Thailand Artinskian 9
Tak Fa Fm - Location O Thailand ? 12
Talung Fm Guangxi Autonomous
Region, China
Late
Changhsingian
Slope and deep basin Bryozoa indet.
Tanquary Fm Ellesmere Island, Canada Asselian Reefs & bioherms Bryozoa indet.
Tarap Fm Tibet Autonomous
Region, China
Norian Outer shelf 2
Tavan-Khara Fm Mongolia Capitanian 11
Tepuel Gp Argentina Asselian 1
Tetyukhinskaya Fm - Dalnegorsky reef
complex
Primor’re region, Russia Norian Reefs & bioherms 2
Thaynes Fm Nevada, United States Olenekian Middle shelf 1
Tianjingshan Fm Sichuan Province, China Ladinian Inner shelf & lagoon Brachiopoda
Togotuiskaya Fm Transbaikal region,
Russia
Capitanian Nearshore 3
Tokrossøya Fm - Bryozoan Horizon Spitzbergen, Norway Late Artinskian-
Kungurian
2
Torlesse Supergroup New Zealand Ladinian Nearshore 1
Torlesse Supergroup New Zealand Ladinian Slope and deep basin 1
Toroweap Fm Nevada, United States Kungurian Inner shelf-Middle shelf 15
139
Lithostratigraphic unit Location Age Environment
Generic
diversity
Trapper Creek Fm Idaho, United States Kungurian Bryozoa indet.
Trogkofel Fm Austria Sakmarian-
Artinskian
Slope mounds 13
Turmiel Marls Fm Spain Toarcian Middle shelf 2
Twillingodden Fm (Sticky Keep Fm) Spitzbergen, Norway Olenekian Middle shelf 1
Ulakhan Fm? Yakutzk Province, Siberia,
Russia
Induan Inner shelf & lagoon 1
unknown formation - Ardeche France Early Toarcian Inner shelf-Middle shelf 1
unknown formation - Balaton Highland Hungary Anisian Inner shelf-Middle shelf Brachiopoda
unknown formation - Calcareous Mbr Urals, Russia Early Roadian Reefs? Bryozoa indet.
unknown formation - Cape Tumul Siberia, Russia Carnian 1
unknown formation - Czech Republic Czech Republic Anisian Inner-Middle-Outer shelf-
Slope
Brachiopoda
unknown formation - East Timor East Timor Norian Reefs & bioherms Bryozoa indet.
unknown formation - Kolyma Basin Siberia, Russia Norian 1
unknown formation - Kulaya River Urals, Russia Wordian 1
unknown formation - N. Caucasus Caucasus region, Georgia Anisian Middle shelf? 1
unknown formation - NW Caucasus Caucasus region, Russia Norian 1
unknown formation - Pamir Tajikistan Artinskian 2
unknown formation - SE Pamirs Tajikistan Carnian-Norian 1
unknown formation - Shapkina Well Urals, Russia Asselian Bryozoa indet.
unknown formation - Sudzukhe River Primor’re region, Russia Ladinian Middle shelf 1
unknown formation - Veszprem beds Hungary Carnian Reefs 4
unknown formation - Volgograd Belt Volgograd, Russia Early Wordian Inner shelf & lagoon 1
unknown formation - Vyatka River Urals, Russia Wordian 1
unknown formation - Western
Kazakhstan
Kazakhstan Early Wordian 1
unnamed formation - Blalock Field Texas, United States Asselian Reefs & bioherms Bryozoa indet.
unnamed formation - Diamond Range Nevada, United States Sakmarian? Outer shelf-Slope and deep
basin
3
unnamed formation - East Cape R. Ellesmere Island, Canada Asselian Reefs & bioherms Bryozoa indet.
unnamed formation - E-Koryak II Kamchatka, Russia Asselian Bryozoa indet.
unnamed formation - Kamishin Volgograd, Russia Asselian Reefs & bioherms Bryozoa indet.
unnamed formation - Karachaganak-
Koblandin Zone
Kazakhstan Asselian Reefs & bioherms Bryozoa indet.
unnamed formation - Kolguev Island Barents Sea, Russia Asselian Reefs & bioherms Bryozoa indet.
140
Lithostratigraphic unit Location Age Environment
Generic
diversity
unnamed formation - Martha van Eman Texas, United States Asselian Slope mounds? Bryozoa indet.
unnamed formation - McKinley Bay Ellesmere Island, Canada Asselian Reefs? Bryozoa indet.
unnamed formation - Urakovs Urals, Russia Asselian-
Sakmarian
Reefs & bioherms Bryozoa indet.
unnamed formation - Vinton Texas, United States Asselian Reefs & bioherms Bryozoa indet.
Upper Bird Spring Fm Nevada, United States Artinskian Bryozoa indet.
Upper Changxing Fm - Laolongdong
reefs
Hubei Province, China Late
Changhsingian
Reefs & bioherms 7
Upper Changxing Fm - Tudiya Buildup Sichuan Province, China Late
Changhsingian
Reefs & bioherms 3
Upper Colina Limestone Mexico Artinskian-
Kungurian
Inner shelf & lagoon Bryozoa indet.
Upper Gechang Fm Himalayas, India Capitanian Nearshore 4
Upper Hanwang Fm Sichuan Province, China Carnian Slope and deep basin Bryozoa indet.
Upper Hanwang Fm Sichuan Province, China Carnian Slope mounds 1
Upper Lugin Gol Fm - Dzhirem-Ula
Mountains
Mongolia Wordian 9
Upper Lugin Gol Fm - Northern Tsagan-
Ula Mountains
Mongolia Wordian Nearshore? 9
Upper Muschelkalk Fm Germany Late Anisian-
Ladinian
Nearshore-Inner shelf-Middle
shelf
Brachiopoda
Upper Pseudoschwagerina Ls Austria Sakmarian Inner shelf & lagoon 5
Upper Serla Fm Italy Late Anisian Reefs & bioherms 1
Utanak Fm Khabarovsk Territory,
Russia
Artinskian 4
Van Hauen Fm Sverdrup basin, Canada Early Kungurian-
Capitanian
Slope and deep basin Bryozoa indet.
Vardebukta Fm - ‘Myalina’ Beds Spitzbergen, Norway Induan-Olenekian Nearshore 1
Vardebukta Fm - Bjornskardet Section Spitzbergen, Norway Induan-Olenekian Outer shelf 1
Verkhnemrujskaia Fm or Kovalskaia
Fm?
Urals, Russia Wordian 3
Vikinghøgda Fm - Deltadalen Mbr Svalbard, Norway Induan Inner shelf & lagoon Brachiopoda
Vladivostok Fm Primor’re region, Russia Roadian-Early
Wordian
Nearshore 23
Vysoka Fm - Geldek Mbr Slovakia Anisian Inner shelf & lagoon 2
141
Lithostratigraphic unit Location Age Environment
Generic
diversity
Wandagee Fm Carnarvon Basin,
Australia
Roadian Outer shelf 5
Wandrawandian Siltstone Sydney Basin, Australia Kungurian Slope and deep basin 10
Wargal Fm - Kalabagh Mbr Salt Range, Pakistan Late Wuchiapingian Inner shelf & lagoon 25
Wargal Fm - Ratti Wahan Mbr Salt Range, Pakistan Capitanian Nearshore Bryozoa indet.
Wargal Fm - Sakesar Mbr Salt Range, Pakistan Wuchiapingian Inner shelf-Middle shelf Bryozoa indet.
Wargal Fm - Topmost Beds Salt Range, Pakistan Changhsingian 1
Wasp Head Fm Sydney Basin, Australia Sakmarian Nearshore 1
Wegener Halvø Fm Greenland, Denmark Late Wuchiapingian Outer shelf - Slope mounds Bryozoa indet.
Werfen Fm - Mazzin Mbr Italy Induan Middle shelf-Outer shelf Brachiopoda
Werfen Fm - Tereso Horizon Italy Late
Changhsingian
Onshore Brachiopoda
West Arm Group Tasmania, Australia Late Artinskian 5
West Arm Group Tasmania, Australia Kungurian 5
West Arm Group Tasmania, Australia Early Roadian 3
Weston Mudstone Tasmania, Australia Late Artinskian-
Kungurian
4
Wetterstein Limestone Fm Slovakia Ladinian Reefs & bioherms Brachiopoda
Whitehorse Sandstone Fm Oklahoma and Texas,
United States
Capitanian Reefs & bioherms 1
Word Fm Texas, United States Wordian Middle shelf-Outer shelf Bryozoa indet.
Wordie Creek Fm Greenland, Denmark Induan Outer shelf Brachiopoda
Wreford Fm - Havensville Shale Mbr Kansas, United States Sakmarian Middle shelf 9
Wreford Fm - Schroyer Limestone Mbr Kansas, United States Sakmarian Middle shelf 15
Wreford Fm - Threemile Limestone Mbr Kansas, United States Sakmarian Nearshore 10
Wujiaping Fm Guangxi Autonomous
Region and Guizhou
Province, China
Wuchiapingian Middle shelf-Outer shelf Bryozoa indet.
Xiala Fm Tibet Autonomous
Region, China
Wordian-
Capitanian
23
Xiala Fm - Units 17 & 18 Tibet Autonomous
Region, China
Wordian-
Capitanian
Brachiopoda
Xiamidong Fm Hunan, Jiangxi, and Hubei
provinces, China
Wuchiapingian Reefs & bioherms Bryozoa indet.
Xiangbo Reef Guangxi Autonomous
Region, China
Wordian-
Capitanian
Reefs & bioherms 7
142
Lithostratigraphic unit Location Age Environment
Generic
diversity
Yamamba Fm Japan Capitanian 5
Yastrebovka Fm Primor’re region, Russia Wuchiapingian Bryozoa indet.
Yates-Tansill Fm - Capitan-Massive Mbr New Mexico, United
States
Capitanian Reefs & bioherms 5
Yenduyet Fm Vietnam Early
Changhsingian
Brachiopoda
Yinkeng Fm Zhejiang Province, China Induan Brachiopoda
Yongde Formation Yunnan Province, China Kungurian-Wordian Outer shelf 8
Zechstein Fm Germany Wuchiapingian Middle shelf 7
Zechstein Fm Germany Wuchiapingian Reefs & bioherms 11
Zewan Fm - Unit A Himalayas, India Capitanian Inner shelf & lagoon? 7
Zewan Fm - Unit B & C Himalayas, India Wuchiapingian Inner shelf & lagoon? 3
Zewan Fm - Unit D Himalayas, India Changhsingian Inner shelf & lagoon 7
Zhamure Fm Tibet Autonomous
Region, China
Carnian Inner shelf & lagoon 2
Zhitkov Fm Primor’re region, Russia Middle Olenekian Outer shelf? Brachiopoda
Zhongbei Melange Tibet Autonomous
Region, China
Wuchiapingian Outer shelf Bryozoa indet.
Zhuravlev strata Komsomolets Island,
Russia
Kungurian 1
Ziyundong Limestone Guizhou Province, China Wordian-
Capitanian
Reefs & bioherms Bryozoa indet.
Zozar Fm Himalayas, India Early Norian Inner shelf & lagoon 3
Zuhánya Limestone Hungary Late Anisian Middle shelf Brachiopoda
143
APPENDIX B
Bryozoan references
Complete list of references used to compile the Permian to Jurassic generic diversity and
distribution (environmental and geographical) of marine stenolaemate bryozoans. The list
also includes references for rhynchonelliform brachiopods.
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144
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190
APPENDIX C
Stratigraphic, environmental, and generic diversity data
Data compiled for Permian to Jurassic marine stenolaemate bryozoans and rhynchonelliform brachiopod (*)
assemblages and used to create Figure 2.1. See Appendix B for detailed references.
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Copacabana Fm
- Section 9
Peru Asselian Middle shelf Cervella, Fenestella, Goniocladia,
Meekopora, Polypora,
Silvaseptopora
Newell et al., 1953
Elephant Canyon
Fm - Bed 20
United States Asselian Inner shelf &
lagoon
Fistulipora, Stenopora, Tabulipora Terrel, 1972
Miharano Fm Japan Asselian Reefs &
bioherms
Anastomopora, Fenestella (Minilya),
Fistulipora, Polypora,
Streblascopora
Niko and Ozawa, 1997
Sakagami and Akagi, 1961
Sakagami. 1970a
Wang et al., 2006
Mt. Bailey Fm Canada Asselian Slope mounds Bryozoa indet. Beauchamp et al, 1991
Beauchamp and Olchowy,
2003
Morin et al., 1994
unknown
formation -
Central Urals
Russia Asselian Outer shelf Fenestrida indet. Izart et al., 1999
191
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Nordenskiölbreen
Fm - Tyrrellfjellet
Mbr
Norway Asselian-
Sakmarian
Nearshore Acanthocladia, Alternifenestella,
Archimedes, Ascopora,
Ascoporella, Coscinum, Eridopora,
Fabifenestella, Goniocladia,
Penniretepora, Polypora, Ramipora,
Rectifenestella, Rhombotrypella,
Streblotrypa, Tabulipora,
Timanodictya
Malecki, 1997
Nakrem, 1988, 1991,
1994a,c
Nakrem et al., 1992
Sunflower Fm United States Asselian-
Sakmarian
Slope and deep
basin
Fistulipora, Penniretepora,
Polypora, Rhabdomeson,
Rhombopora, Rhombotrypella,
Sulcoretepora
Coash, 1967
Schwarz et al, 1991
Stuart, 1962
Callytharra Fm Australia Sakmarian Inner shelf &
lagoon
Dyscritella, Evactinopora,
Fenestella, Hexagonella,
Linguloclema, Lyropora,
Penniretepora, Polypora,
Protoretepora, Ramipora,
Rhabdomeson, Rhombocladia,
Rhombopora?, Saffordotaxis,
Septopora, Streblocladia,
Streblotrypa (Streblascopora)
Dixon, 2002
Engel, 1987
Eyles et al., 2003
Ross, 1963b
Nelynya-shor Fm Russia Sakmarian Outer shelf Anisotrypella, Paraseptopora,
Uralotrypa
Belyakov, 1994
Lisitsyn, 1991
Trogkofel Fm Austria Sakmarian Slope mounds Alternifenestella, Carnocladia,
Eridopora, Goniocladia,
Paraptylopora, Penniretepora,
Polypora, Primorella, Prismopora,
Rhabdomeson, Rhombopora,
Stenophragmidium, Streblascopora
Buttersack and
Boeckelmann, 1984
Ernst, 2001a, 2003
Flügel, 1981a
Krainer and Davydov, 1988
Samankassou, 2003
192
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Wreford Fm -
Schroyer
Limestone Mbr
United States Sakmarian Middle shelf Acanthocladia, Fenestella,
Fistulipora, Meekopora, Minilya,
Penniretepora, Polypora,
Protoretepora, Rhombopora,
Septopora, Syringoclemis,
Tabulipora
Pachut and Cuffey, 1991,
1999
Newton, 1971
Simonsen, 1977
Akiyoshi
Limestone Gp
Japan Sakmarian-
Artinskian
Reefs &
bioherms
Fistulipora, Meekoporella,
Stenopora, Streblascopora
Ota, 1977
Sakagami and Sugimura,
1981
Sugimura and Ota, 1980
Bulgadoo Shale Australia Artinskian Slope and deep
basin
Hexagonella Moore & Hocking, 1983
Skwarko, 1993a
Cathedral
Mountain Fm -
Poplar Tank Mbr
United States Artinskian Slope mounds Acanthocladia, Fistulipora?,
Penniretepora, Polypora,
Polyporella, Protoretepora,
Rhabdomeson?, Rhombopora,
Septopora, Streblascopora,
Synocladia, Tabulipora,
Timanodictya? Ulrichotrypa?
Ross and Ross, 2003
Zimmerman and Cuffey,
1985, 1987
Cattle Creek Fm Australia Artinskian Inner shelf &
lagoon
Diploporaria, Fenestella, Polypora,
Polyporella, Protoretepora
Fielding, 1989
Fielding et al., 2000
Jackson et al., 1980
Jones et al., 2006
Palmieri, 1998
Wass, 1968a
193
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Great Bear Cape
Fm
Canada Artinskian Middle shelf Acanthocladia, Alternifenestella,
Bashkirella, Clausotrypa,
Cyclotrypa, Dyscritella,
Fabifenestella, Fenestella,
Fistulipora, Penniretepora,
Phragmophera, Polypora,
Primorella, Pseudonematopora,
Ramipora, Rectifenestella,
Rhombopora, Rhombotrypella,
Stenophragmidium, Streblascopora,
Tabulipora, Ulrichotrypa
Beauchamp and Henderson,
1994
Ernst and Nakrem, 2005
Kells et al., 1995
Rat Buri Ls - Ko
Muk Island
Thailand Artinskian Outer shelf Acanthocladia, Ascopora,
Dyscritella, Fenestella, Fenestella
(Minilya), Fistulipora, Goniocladia,
Penniretepora, Polypora,
Rhabdomeson, Streblascopora,
Streblotrypa, Sulcoretepora
Fontaine et al., 1994
Grant, 1976
Sakagami, 1966a, b
Sakagami, 1970a, b
Kapp Starostin
Fm - Vøringen
Mbr
Norway Artinskian Nearshore Alternifenestella, Clausotrypa,
Cyclotrypa, Dyscritella,
Fabifenestella, Fenestella,
Girtyporina, Goniocladia, Lyropora,
Meekopora, Penniretepora,
Permoheloclema, Polypora,
Polyporella, Primorella, Ramipora,
Rectifenestella, Rhombotrypella,
Stenopora, Streblascopora,
Tabulipora, Timanodictya
Malecki, 1977
Nakrem, 1998, 1991
Nakrem, 1994a, b
Nakrem et al., 1992
Balikelik Fm China Kungurian Inner shelf &
lagoon
Ascopora, Dybowskiella,
Eridopora?, Fenestella, Fistulipora,
Klaucena (Klaucena), Maychella,
Penniretepora, Rhabdomeson,
Rhombopora, Streblascopora,
Streblotrypa, synocladia, Tabulipora
Ruan, 1996a
Yang and Lu, 1983
194
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Cathedral
Mountain Fm -
Wedin Congl.
Mbr
United States Kungurian Slope mounds Acanthocladia, Anastomopora,
Fenestella, Penniretepora,
Polypora, Rhombopora?,
Septopora, Timanodictya?
Ross and Ross, 2003
Zimmerman and Cuffey,
1985, 1987
Kozhim Fm Russia Kungurian Middle shelf Cyclotrypa, Dyscritella,
Neoeridotrypella
Kotlyar, 2002
Lisitsyn, 1986
Miseryfjellet Fm Norway Kungurian Nearshore Anisotrypella, Dyscritella,
Dyscritellina, Fenestella,
Gilmouropora, Kingopora,
Lyrocladia, Permoheloclema,
Polypora, Polyporella, Ramipora,
Rectifenestella, Reteporidra,
Rhombotrypella, Septopora,
Stenopora, Tabulipora,
Timanodictya, Wjatkella
Malecki, 1977
Nakrem, 1988, 1991, 2005
Nakrem et al., 1992
Nooncanbah Fm Australia Kungurian Outer shelf Australopolypora, Crockfordia,
Dybowskiella, Dyscritella,
Dyscritellina?, Eridopora, Etherella,
Evactinostella, Fenestella,
Fistulamina, Fistulipora,
Goniocladia, Hexagonella,
Hinganella?, Liguloclema,
Megacanthopora, Minilya,
Ogbinopora?, Polypora, Polyporella,
Prismopora, Protoretepora,
Pseudobatostomella, Ramipora,
Rhabdomeson, Rhombocladia,
Rhombopora?, Saffordotaxis,
Stenodiscus, Stenopora,
Streblotrypa, Synocladia,
Tabulipora
Crockford, 1944
Engel, 1987
Skwarko, 1993a
Stratigraphic Units Database
(Australian Government
Geoscience Australia)
195
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Wandrawandian
Siltstone
Australia Kungurian Slope and deep
basin
Dyscritella, Fenestella,
Laxifenestella, Levifenestella,
Minilya, Paucipora, Polypora,
Rectifenestella, Shulgapora,
Stenopora
Eyles et al., 1998
Ramli and Crook, 1978
Reid, 2003
Tye et al., 1996
Road Canyon Fm United States Kungurian-
Roadian
Reefs &
bioherms
Acanthocladia, Acanthoclema,
Anastomopora, Fenestella,
Fistulipora?, Isotrypa,
Parafenestralia, Penniretepora,
Polypora, Protoretepora,
Rhabdomeson, Rhombopora?,
Septopora, Stenodiscus?,
Stenopora?, Synocladia,
Tabulipora, Timanodictya?,
Ulrichotrypa?
Ross and Ross, 2003
Zimmerman and Cuffey,
1985, 1987
Harries et al., 2000
Iwaizaki Fm -
Unit 4
Japan Roadian Nearshore Fenestella, Fistulipora,
Hayasakapora, Meekopora,
Pseudobatostomella, Saffordotaxis,
Streblascopora, Sulcoretepora
Kawamura and Machiyama,
1995
Sakagami, 1961, 1970a,
1973b
Shen and Kawamura, 2001
Kaibab Fm - Beta
Mbr
United States Roadian Inner shelf &
lagoon
Batostomella, Bicorbis, Cystodictya,
Fenestella, Fistulipora, Girtypora,
Leioclema, Leptopora, Meekopora,
Polypora, Protoretepora?,
Rhabdomeson, Rhobopora,
Septopora, Stenodiscus?,
Stenopora, Streblotrypa
Chronic, 1952
McKinney, 1983
Kapp Starostin
Fm
Norway Roadian Outer shelf Alternifenestella, Clausotrypa,
Dyscritella, Fabifenestella,
Lyrocladia, Permoheloclema,
Polypora, Primorella, Ramipora,
Rectifenestella, Stenopora
Malecki, 1977
Nakrem, 1988, 1991, 1994a,
2005
Nakrem et al., 1992
196
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Kapp Starostin
Fm
Norway Roadian-
Wordian
Middle shelf Anisotrypella, Cyclotrypa,
Dyscritella, Fistulipora,
Gilmouropora, Neoeridotrypella,
Polypora, Reteporidra,
Rhombotrypella, Polypora,
Reteporidra, Rhombotrypella,
Stenopora, Tabulipora,
Timanodictya
Malecki, 1977
Nakrem, 1988, 1991, 1994a,
2005
Nakrem et al., 1992
Van Hauen Fm Canada Roadian-
Capitanian
Slope and deep
basin
Bryozoa indet. Beauchamp and Lamirande,
1990
Desrochers and Beauchamp,
1995
Kanokura Fm Japan Wordian Inner shelf &
lagoon
Fenestella, Fistulipora, Leioclema,
Meekopora, Penniretepora,
Polypora, Pseudobatostomella,
Septopora, Sulcoretepora,
Tabulipora?, Thamniscus
Kawamura and Machiyama,
1995
Minato et al., 1978
Sakagami, 1961, 1970a
Shi, 2006
Mission Argillite
Fm
United States Wordian Reefs &
bioherms
Alternifenestella, Coeloclemis,
Dybowskiella, Dyscritella,
Dyscritellina, Fistulamina?,
Fistulipora, Fistuliramus,
Hayasakapora, Mackinneyella,
Meekoporella, Nematopora,
Neoeridotrypella, Pamirella,
Parapolypora, Pinegopora,
Polypora, Polyporella,
Pseudobatostomella,
Rhombopora?, Rhombotrypella,
Sakagamiina, Stenopora,
Streblotrypa?, Tabulipora, Wjatkella
Gilmour and Snyder, 2000
Murdock
Mountain Fm
United States Wordian Outer shelf Dyscritella, Hinganella, Kingopora,
Morozoviella, Neoeridotrypella,
Pseudobatostomella, Stenodiscus,
Stenopora, Streblotrypa,
Thamniscus
Gilmour et al., 1997
197
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Pietra di
Salomone
Megablock
Italy Wordian Slope mounds Fistulipora, Hayasakapora Ernst, 2000
Flügel et al., 1991
Sokyuiskaya Fm Russia Wordian Nearshore Dyscritella, Maychella, Maychellina,
Permofenestella, Stenopora,
Streblascopora
Kotlyar et al., 2006
Akiyoshi
Limestone Gp
Japan Capitanian Inner shelf &
lagoon
Clausotrypa, Dyscritella, Eridopora,
Fistulipora, Girtyporina,
Hayasakapora, Hexagonella,
Paralioclema, Permolioclema,
Pseudobatostomella,
Rhabdomeson, Rhombopora,
Streblascopora
Sakagami and Sugimura,
1981, 2000
Capitan Fm United States Capitanian Slope mounds Fenestella, Goniocladia, Polypora Fagerstrom and Weidlich,
2005
Wood, 1999
Chandalaz Fm Russia Capitanian Nearshore Cyclotrypa, Dyscritellina,
Epiactinotrypa, Etherella,
Fistulipora, Girtypora,
Hayasakapora, Hinganella,
Neoeridocampylus, Ogbinopora,
Parastenodiscus, Prismopora,
Tavayzopora, Ulrichotrypella
Belyaeva et al., 1997
Kiseleva, 1982b
Kotlyar et al., 2006
Gerster
Limestone
United States Capitanian Middle shelf Araxopora, Cyclotrypa, Dyscritella,
Dyscritellina, Fenestella, Fistulipora,
Hinganella, Kingopora, Meekopora,
Morozoviella, Neoeridotrypella,
Paralioclema, Polypora, Polyporella,
Pseudobatostomella,
Rectifenestella, Reteporidra,
Rhombotrypella, Stellahexiformis,
Stenodiscus, Stenopora,
Timanodictya, Timanotrypa,
Ulrichotrypella, Wjatkella
Gilmour and Snyder, 1986
Snyder, 1976
Snyder and Gilmour. 2006
Wardlaw, 1974
198
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Xiangbo reef China Capitanian Reefs &
bioherms
Ascopora, Dybowskiella, Eridopora,
Fenestella, Fistulipora, Polypora,
Rhabdomeson
Sheng, 1991
Hambast Fm Iran Wuchiapingian Outer shelf Fistulipora, Polypora Heydary et al., 2003
Sakagami, 1980
Taraz, 1971
Taraz et al., 1981
Qubuerga Fm China (Tibet) Wuchiapingian Nearshore Araxopora?, Dyscritella,
Fistuliramus, Maychella, Polypora,
Rhombopora, Stenopora
Sakagami et al., 2006
Shen et al. 2001
Shen et al., 2006
Zhang, 1987
Saser Brangsa
Fm*
India Wuchiapingian Slope and deep
basin
Shinha et al., 1999
Waterhouse and Gupta,
1983
Wargal Fm -
Kalabagh Mbr
Pakistan Wuchiapingian Inner shelf &
lagoon
Acanthocladia, Alternifenestella,
Dybowskiella, Dyscritella,
Fabifenestella, Fistulamina,
Fistulipora, Hayasakapora,
Hexagonella, Laxifenestella,
Mackinneyela, Minilya, Montrypa,
Nematopora, Parapolypora,
Penniretepora, Polypora,
Polyporella, Pseudobatostomella,
Rectifenestella, Rhabdomeson,
Rhombopora, Stenopora,
Streblascopora, Streblotrypa
Ali et al., 1985
Gilmour and Morozova, 1999
Mertmann, 2003
Wegener Halvø
Fm
Greenland Wuchiapingian Slope mounds Fenestrida indet., Trepostomida
indet.
Hurst et al., 1989
Nielsen and Shen, 2004
Stemmerik, 2001
Zechstein Fm Germany Wuchiapingian Reefs &
bioherms
Acanthocladia, Dyscritella,
Fenestella, Kingopora,
Penniretepora, Rectifenestella,
Ryhopora, Spinofenestella,
Stenopora, Synocladia, Thamniscus
Ernst, 2001a
Lisitsyn and Ernst, 2004
199
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Zechstein Fm Germany Wuchiapingian Middle shelf Dyscritella, Kavariella, Kingopora,
Penniretepora, Spinofenestella,
Thamniscus, Ulrichotrypa
Ernst, 2001a
Lisitsyn and Ernst, 2004
Changhsing Fm China Changhsingian Middle shelf Araxopora, Dybowskiella,
Fenestella, Fistulipora, Polypora,
Protoretepora, Stenopora
Chen et al., 2005
Fan et al., 1990
Fan et al., 1982
Yang and Lu, 1980, 1981
Upper
Changhsing Fm
China
Changhsingian
Reefs &
bioherms
Acanthocladia, Clausotrypa,
Dybowskiella, Fenestella,
Fistulipora, Polypora, Sulcoretepora
Fan et al., 1990
Fan et al., 1982
Yang and Lu, 1981
Chhidru Fm Pakistan Changhsingian Nearshore Acanthocladia, Dybowskiella,
Dyscritella, Hexagonella,
Mackinneyella, Monotrypa,
Parapolypora, Protoretepora,
Septopora, Stenodiscus, Stenopora,
Synocladia, Thamniscus
Gilmour and Morozova, 1999
Kapoor and Tokuoka, 1985
Mertmann, 2003
Episkopi Fm Greece Changhsingian Outer shelf Acanthocladia Clapham, 2006
Talung Fm China Changhsingian Slope and deep
basin
Bryozoa indet. He et al., 2005
Zewan Fm India Changhsingian Inner shelf &
lagoon
Diploporaria, Dyscritella, Fistulipora,
Hayasakapora, Septopora,
Stenodiscus, Stenopora
Chen et al., 2005
Kapoor and Nakazawa, 1981
Sakagami, 1981
Shen et al., 2006
Tewari and Srivastava, 1967
Dinwoody Fm* United States Induan Middle shelf Rodland and Bottjer, 2001
Feixianguan Fm -
Member 1*
China Induan Inner shelf &
lagoon
Reinhardt, 1988
Kistefjellet Fm Norway Induan Slope and deep
basin
Paralioclema Nakrem and Mørk, 1991
Worsley and Mørk, 1978
200
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Vardebukta Fm -
‘Myalina’ Beds
Norway Induan-
Olenekian
Nearshore Paralioclema Mørk et al., 1982
Nakrem and Mørk, 1991
Wignall et al., 1998
Worsley and Mørk, 1978
Vardebukta Fm -
Bjornskardet
Section
Norway Induan-
Olenekian
Outer shelf Paralioclema Mørk et al., 1982
Nakrem and Mørk, 1991
Wignall et al., 1998
Worsley and Mørk, 1978
Blind Fjord Fm Canada Olenekian Slope and deep
basin
Arcticopora Bolton, 1962
Devaney, 1991
Fritz, 1962
Moenkopi Fm -
Virgin Limestone
Mbr*
United States Olenekian Inner shelf &
lagoon
Fraiser, 2005
Thaynes Fm United States Olenekian Middle shelf Dyscritellopsis Schäfer et al., 2003
Dixie Valley Fm United States Anisian Nearshore Phragmotrypa Carey, 1984
Nichols and Siberling, 1977
Schäfer and Fois, 1987
Dont Fm Italy Anisian Reefs &
bioherms
Reptonoditrypa Schäfer, 1994
Schäfer and Fois, 1987
Senowbari-Daryan et al.,
1993
Favret Fm United States Anisian Middle shelf Phragmotrypa Carey, 1984
Nichols and Siberling, 1977
Schäfer and Fois, 1987
Karazin Fm Russia Anisian Outer shelf Paralioclema Burij, 1997
Burij et al., 1993
Morozova, 1969
Schäfer, 1994
Zakharov et al., 2000
Schlern Fm Italy Anisian Slope and deep
basin
Reptonoditrypa Emmerich et al., 2005
201
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Vysoká Fm -
Geldek Mbr
Slovakia Anisian Inner shelf &
lagoon
Dyscritella?, Vysokella Michalík et al., 1992
Zágorsek, 1993
Laibuxi Fm China (Tibet) Ladinian Inner shelf &
lagoon
Paralioclema, Pseudobatostomella,
Tebitopora
Hu, 1984
Liu and Xu, 1998
Schäfer, 1994
Yang and Xia, 1975
Liard Fm Canada Ladinian Reefs &
bioherms
Bryozoa indet. Zonneveld, 2001
Torlesse
Supergroup
New Zealand Ladinian Nearshore Dyscritellopsis
Schäfer and Grant-Mackie,
1994
Torlesse
Supergroup
New Zealand Ladinian Slope and deep
basin
Pseudobatostomella Schäfer and Grant-Mackie,
1994
unknown
formation -
Sudzukhe River
Russia Ladinian Middle shelf Pseudobatostomella Morozova, 1969
Mukut Fm* Nepal Ladinian-
Carnian
Outer shelf Fuchs et al., 1988
Garzanti, 1999
Siblík, 1975
Cassian Fm* Italy Carnian Slope and deep
basin
Fürsich and Wendt, 1977
Naizawa Fm Japan Carnian Middle shelf Dyscritella, Leioclema,
Pseudobatostomella, Zozariella
Sakagami and Sakai, 1976
Schäfer, 1994
Oxytoma-Mytalus
beds
Japan Carnian Nearshore Pseudobatostomella Ando, 1987
Sakagami, 1972
San Cassiano
Fm
Italy Carnian Slope mounds Braiesopora, Corynotrypoides,
Cystitrypa, Dyscritella, Stomatopora
Bizzarini and Braga, 1978,
1981, 1985, 1993, 1994
Schäfer, 1994
Schäfer and Fois, 1987
unknown
formation -
Veszprem beds
Hungary Carnian Reefs &
bioherms
Dyscritella, Monotrypa, Stenopora,
Stomatopora
Bizzarini and Braga, 1982
Morozova, 1969
202
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Zhamure Fm China (Tibet) Carnian Inner shelf &
lagoon
Dyscritella, Paralioclema Liu and Xu, 1998
Yang and Xia, 1975
Yin et al., 1987
Luning Fm* United States Norian Middle shelf Hogler, 1992
Sandy, 1995
Sandy and Stanley, 1993
Murihiku Terrane New Zealand Norian Nearshore Dyscritella, Stenodiscus Schäfer and Grant-Mackie,
1994
Sotong
Limestone
Malaysia Norian Slope mounds Bryozoa indet. Fontaine et al., 1990
Tarap Fm China (Tibet) Norian Outer shelf Tebitopora, Zozarellia Garzanti, 1999
Jadoul et al., 1998
Sagakami et al., 2006
Tetyukhinskaya
Fm
Russia Norian Reefs &
bioherms
Buria, Reptonodicava Buryi, 1988
Morozova and Zharnikova,
1984
Punina, 1997
Zozar Fm India Norian Inner shelf &
lagoon
Dycritella, Tebitopora, Zozariella Baud et al., 1984
Gaetani and Garzanti, 1991
Gaetani et al., 1986
Hu, 1984
Schäfer and Fois, 1987
Kössen Fm* Austria Norian-
Rhaetian
Slope and deep
basin
Golebiowski, 1989
Piller, 1981
Sandy, 1995
Turnsek et al., 1999
Fatra Fm* Slovakia Rhaetian Inner shelf &
lagoon
Tomašových, 2004
Hybe Fm Slovakia Rhaetian Middle shelf Reptomultisparsa, Stomatopora Gazdzizki, 1974
Taylor and Michalík, 1991
203
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Nayband Fm Iran Rhaetian Reefs &
bioherms
Cyclotrypa, Diaphragmopora,
Dyscritella, Reptonoditrypa
Schäfer et al., 2003
Redcar
Mudstone Fm
England Hettangian Middle shelf Stomatopora Cox et al., 1999
Taylor, 2006
Kendlbach Fm* Austria Hettangian Inner shelf &
lagoon
Leopold et al., 2005
Siblík, 1999
Schnoll Fm* Austria Hettangian Slope and deep
basin
Boehm et al., 1999
Leopold et al., 2005
Blue Lias Fm -
Porthkerry Mbr
England Sinemurian Middle shelf Oncousoecia Sheppard et al., 2006
Simms et al., 2002
Taylor, 2006
Warrington and Ivimey-Cook,
1995
Calcari Grigi Fm Italy Sinemurian Inner shelf &
lagoon
Bryozoa indet. Broglio Loriga et al., 1991
Fraser et al., 2004
Amaltheen Fm Germany Pliensbachian Middle shelf-
outer shelf
Berenicea, Proboscina,
Stomatopora
Illies, 1971, 1973
Jablonski et al., 1997
Taylor, 2006
Dyrham Fm England Pliensbachian Middle shelf-
outer shelf
Berenicea, Reptomultisparsa Cox et al., 1999
Taylor, 2006
Marlstone Rock
Fm
England Pliensbachian Outer shelf Mesenteripora, Spiropora Hallam, 1967
Tate, 1875
Taylor, 2006
Taylor and Sequeiros, 1982
Walford, 1887
Marlstone Rock
Fm
England Toarcian Outer shelf Mesenteripora, Stomatopora Hallam, 1967
Tate, 1875
Taylor, 2006
Taylor and Sequeiros, 1982
Walford, 1887
S. Gião Unit* Portugal Toarcian Slope mounds Duarte et al., 2001
204
Lithostratigraphic
unit
Country Age Environment Bryozoan genera References
Turmiel Marls Fm Spain Toarcian Middle shelf Microeciella , ‘Proboscina’ Fürsich et al., 2001
Perrili, 2000
Taylor and Sequeiros, 1982
unknown
formation -
Ardèche
France Toarcian Inner shelf &
lagoon
Radicipora Jablonski et al., 1997
Walter, 1970
205
APPENDIX D
Bryozoan stage-level generic diversity
Permian to Early Jurassic stage-level presence data (indicated with an X) for all stenolaemate bryozoans. Abbreviated
stages for the Permian: Wuchiap. = Wuchiapingian, Chang. = Changhsingian. Abbreviated stages for the Jurassic:
Hettan. = Hettangian, Sinem. = Sinemurian, Pliens = Pliensbachian, Toarc. = Toarcian, Aalen. = Aalenian, Bajoc. =
Bajocian, Bathon. = Bathonian, Callov. = Callovian, Oxfor. = Oxfordian, Kimmer. = Kimmeridgian, Tithon. =
Tithonian. References used are listed in Appendix B.
PERMIAN
Asselian Sakmarian Artinskian Kungurian Roadian Wordian Capitanian Wuchiap. Chang.
Order
Cryptostomida
Acanthoclema ? ?
Ascopora X X X X - X
Ascoporamagna ? ?
Ascoporella X X
Clausotrypa X - X X - X X - X
Cystodictya ? ?
Dichotrypa X - X ?
Gilmouropora X X ? ?
Girtypora X X X
Girtyporina X X X X X X X
Hayasakapora X X X X X X X X X
Hyphasmopora X
Klaucena ? ? ? ?
Maychella ? X X X X ?
206
PERMIAN
Asselian Sakmarian Artinskian Kungurian Roadian Wordian Capitanian Wuchiap. Chang.
Maychellina X X X
Megacanthopora X
Morozoviella X X
Nematopora X - X
Nicklesopora X - - - X
Nikiforovella X X X
Ogbinopora X - X X
Paleschara X
Pamirella X X - - X X
Permoheloclema X X ? ?
Phragmophera X
Pictatella X X
Pinegopora X X
Primorella X - X X - X X
Pseudonematopora X
Rhabdomeson X X X X X X X X X
Rhombopora X X X X X X X X X
Rhomboporella ? ?
Robinella X
Saffordotaxis X X X X - X
Sakagamiina X
Streblascopora X X X X X X X X X
Streblocladia X
Streblotrypa X X X X X X X X
Syringoclemis X - X ? ?
Tavayzopora X X
Tebitopora
Timanodictya X X X X X X X
Timanotrypa X - X
Uralotrypa X
207
PERMIAN
Asselian Sakmarian Artinskian Kungurian Roadian Wordian Capitanian Wuchiap. Chang.
Order
Cyclostomida
Apsendesia
Bere
Braiesopora
Buria
Cava
Ceriocava
Ceriopora
Collapora
Corynotrypa X - - - - - - X
Corynotrypoides
Crescis
Diastopora
Entalopora
Fasciculipora
Hederella X
Heteropora
Hyporosopora
Idmonea
Kololophos
Leptopora X
Mecynoecia
Mesenteripora
Mesonopora
Microeciella
Multisparsa
Multitubigera
Oncousoecia
Patulopora
Plagioecia
Proboscinopora
208
PERMIAN
Asselian Sakmarian Artinskian Kungurian Roadian Wordian Capitanian Wuchiap. Chang.
Radicipora
Reptoclausa
Reptomultisparsa
Reptonodicava
Reticulipora
Ripisoecia
Spiropora
Stomaporina
Stomatopora
Stomatoporopsis
Terebellaria
Tetrapora
Theonoa
Unitubigera
Order Cystoporida
Actinotrypella X
Cassianopora
Coelocaulis ? ?
Coscinium X X
Coscinotrypa X - - - - X X
Cyclotrypa X X X X X - -
Cystitrypa
Dybowskiella X - X X X X
Epiactinotrypa X X
Eridopora X X X X - X X X X
Etherella X X X X
Evactinopora X
Evactinostella X X - X
Filiramoporina X
Fistulamina X - X X X
Fistulipora X X X X X X X X X
209
PERMIAN
Asselian Sakmarian Artinskian Kungurian Roadian Wordian Capitanian Wuchiap. Chang.
Fistuliramus X - X - - - X
Fistulotrypa X ?
Goniocladia X X X X X X X
Hexagonella X X X X X X X X
Liguloclema X X X
Meekopora X X X X X X X
Meekoporella X X - X
Prismopora X X X - X X
Ramipora X X X X - X X
Ramiporidra X X X X - X
Sulcoretepora X - X X X X X - X
Volgia ? ?
Order Fenestrida
Acanthocladia X X X X X - X X X
Alternifenestella X X X X - X - X
Anastomopora X - - X ?
Archimedes X X X - - X
Australopolypora X
Bashkirella X
Bicorbis X
Carnocladia X ?
Cavernella X
Cervella X X
Chainodictyon X
Diploporaria X - - - - X X
Dogaddenella X ?
Exfenestella X - - - X
Fabifenestella X X X X - X X X X
Fenestella X X X X X X X X X
Fenestrellina X
Flexifenestella X X - X ? ?
210
PERMIAN
Asselian Sakmarian Artinskian Kungurian Roadian Wordian Capitanian Wuchiap. Chang.
Hinganotrypa X - X X
Isotrypa ? ?
Kalvariella X - - X X
Kingopora X X X X X X
Laxifenestella X - X X X - - X
Levifenestella X X X
Lyrocladia X - X X ? ?
Lyropora X X X
Mackinneyella X X - X X X X X
Minilya X X X X X X - X
Ogbinofenestella X
Parafenestralia ? X
Parapolypora X X - X X X X X
Paraptylopora ? ?
Paraseptopora X - X
Paucipora X X - - X
Penniretepora X X X X - X X X
Permofenestella X X X
Polypora X X X X X X X X X
Polyporella X X X X - X X X
Protoretepora X X X X X - X X
Pseudopolypora ?
Ptylopora X - X X - - - X
Rectifenestella X X X X X X X X
Reteporidra X - X X X X X
Rhombocladia X X X
Ryhopora X
Septopora X X X X X X X X X
Shulgapora X X
Silvaseptopora X X
Spinofenestella X X X X X X X X
Synocladia X - X X - X X X X
211
PERMIAN
Asselian Sakmarian Artinskian Kungurian Roadian Wordian Capitanian Wuchiap. Chang.
Synocladiella X
Thamniscus X - - X - X - X X
Triznella X
Wjatkella X X X X X X
Order
Trepostomida
Amphiporella - X X
Anisotrypella X - X - X - X
Araxopora X - X X X X
Arcticopora X X
Batostomella X - - X
Callocladia ? ?
Ceoloclemis X - X
Crockfordia X X
Diaphragmopora
Dyscritella X X X X X X X X X
Dyscritellina X X X X X X
Dyscritellopsis
Hinaclema X X X
Hinganella X - X X
Hyalotoechus X - X
Iraidina X X
Leioclema X X X X X X - -
Metelipora X
Monotrypa X X
Neoeridocampylus X
Neoeridotrypella X X X X X
Orbipora X
Paralioclema X - X X X X X
Paramaychella X
Parastenodiscus X X
212
PERMIAN
Asselian Sakmarian Artinskian Kungurian Roadian Wordian Capitanian Wuchiap. Chang.
Permolioclema X
Permopora ? ?
Phragmotrypa
Pseudobatostomella X - X X X X X X
Pycnopora X - X
Reptonoditrypa
Rhombotrypella X X X X X X X
Ruzhencevia X X X
Stellahexiformis X
Stenodiscus X - X X X X X X
Stenophragmidium X X X
Stenopora X X X X X X X X X
Styloclema
Tabulipora X X X X X X X - X
Ulrichotrypa X X - - - X
Ulrichotrypella X X X
Vysokella
Zozariella
213
TRIASSIC
Induan Olenekian Anisian Ladinian Carnian Norian Rhaetian
Order Cryptostomida
Acanthoclema
Ascopora
Ascoporamagna
Ascoporella
Clausotrypa
Cystodictya
Dichotrypa
Gilmouropora
Girtypora
Girtyporina
Hayasakapora
Hyphasmopora
Klaucena
Maychella
Maychellina
Megacanthopora
Morozoviella
Nematopora
Nicklesopora
Nikiforovella
Ogbinopora
Paleschara
Pamirella
Permoheloclema
Phragmophera
Pictatella
Pinegopora
Primorella
Pseudonematopora
Rhabdomeson
Rhombopora
214
TRIASSIC
Induan Olenekian Anisian Ladinian Carnian Norian Rhaetian
Rhomboporella
Robinella
Saffordotaxis
Sakagamiina
Streblascopora
Streblocladia
Streblotrypa
Syringoclemis
Tavayzopora
Tebitopora X - X
Timanodictya
Timanotrypa
Uralotrypa
Order Cyclostomida
Apsendesia
Bere
Braiesopora X
Buria X
Cava
Ceriocava
Ceriopora
Collapora
Corynotrypa
Corynotrypoides X
Crescis
Diastopora
Entalopora
Fasciculipora
Hederella
Heteropora
Hyporosopora
215
TRIASSIC
Induan Olenekian Anisian Ladinian Carnian Norian Rhaetian
Idmonea
Kololophos
Leptopora
Mecynoecia
Mesenteripora
Mesonopora
Microeciella
Multisparsa
Multitubigera
Oncousoecia
Patulopora
Plagioecia
Proboscinopora
Radicipora
Reptoclausa
Reptomultisparsa X
Reptonodicava X
Reticulipora
Ripisoecia
Spiropora
Stomaporina
Stomatopora X - X
Stomatoporopsis
Terebellaria
Tetrapora
Theonoa
Unitubigera
Order Cystoporida
Actinotrypella
Cassianopora X
Coelocaulis
216
TRIASSIC
Induan Olenekian Anisian Ladinian Carnian Norian Rhaetian
Coscinium
Coscinotrypa
Cyclotrypa - - - - - - X
Cystitrypa X
Dybowskiella
Epiactinotrypa
Eridopora
Etherella
Evactinopora
Evactinostella
Filiramoporina
Fistulamina
Fistulipora
Fistuliramus
Fistulotrypa
Goniocladia
Hexagonella
Liguloclema
Meekopora
Meekoporella
Prismopora
Ramipora
Ramiporidra
Sulcoretepora
Volgia
Order Trepostomida
Amphiporella
Anisotrypella
Araxopora
Arcticopora - X
Batostomella
217
TRIASSIC
Induan Olenekian Anisian Ladinian Carnian Norian Rhaetian
Callocladia
Ceoloclemis
Crockfordia
Diaphragmopora X
Dyscritella - - X X X X X
Dyscritellina
Dyscritellopsis X - X X X
Hinaclema
Hinganella
Hyalotoechus
Iraidina
Leioclema - - - - X
Metelipora
Monotrypa - - - - X
Neoeridocampylus
Neoeridotrypella
Orbipora
Paralioclema X X X X X X
Paramaychella
Parastenodiscus
Permolioclema
Permopora
Phragmotrypa X
Pseudobatostomella X - X X X X
Pycnopora
Reptonoditrypa X - - - X
Rhombotrypella
Ruzhencevia
Stellahexiformis
Stenodiscus - - - - - X
Stenophragmidium
Stenopora - - - - X
218
TRIASSIC
Induan Olenekian Anisian Ladinian Carnian Norian Rhaetian
Styloclema X
Tabulipora
Ulrichotrypa
Ulrichotrypella
Vysokella X
Zozariella X X
219
JURASSIC
Hettan. Sinem. Pliens. Toarc. Aalen. Bajoc. Bathon. Callov. Oxfor. Kimm. Tithon.
Order Cyclostomida
Apsendesia X X X X X
Bere
Braiesopora
Buria
Cava X X X
Ceriocava X X X X X
Ceriopora X X X
Collapora X X X X
Corynotrypa
Corynotrypoides
Crescis X X X
Diastopora X X X X X
Entalopora X X X
Fasciculipora X
Hederella
Heteropora X X X X
Hyporosopora X X X X - X
Idmonea X X X X X
Kololophos X X X
Leptopora
Mecynoecia X X X X X
Mesenteripora X X X X X X X - X
Mesonopora X X X X
Microeciella X X - - X
Multisparsa X X X X X
Multitubigera X
Oncousoecia X - - - X X X X X X
Patulopora X
Plagioecia X
Proboscinopora X X
Radicipora X - X X - X X
220
JURASSIC
Hettan. Sinem. Pliens. Toarc. Aalen. Bajoc. Bathon. Callov. Oxfor. Kimm. Tithon.
Reptoclausa X - - - - - X
Reptomultisparsa - - X - X X X X X
Reptonodicava
Reticulipora X X
Ripisoecia X X X X X
Spiropora X X X
Stomaporina X
Stomatopora X X X X X X X X X X X
Stomatoporopsis X X X
Terebellaria X X X X
Tetrapora X
Theonoa X X X X X
Unitubigera X
221
APPENDIX E
Bryozoan colonial morphology
Permian to Early Jurassic morphological data for colonies of stenolaemate bryozoan species. Species are arranged
according to age and formation. Environmental data for each of the assemblages are available in Appendix A.
References used are listed in Appendix B.
Age Formation Genus Species Morphology
Asselian Polypora lyndoni Erect
Stenopora dickinsi Ramose
Stenopora fisheri Ramose
Lyons Group
Stenopora lyndoni Ramose
Asselian Sulcoretepora corticata Bifoliate
Sulcoretepora irregularis Bifoliate frond
Rhabdomeson sp. Cylindrical
Fistulipora bifoliatus Erect Bifoliate
Polypora sp. Fenestrate
Fistulipora lamellatis Lamellate
Penniretepora sp. Pinnate
Rhombopora cornwallis Ramose
Sunflower Fm
Rhombotrypella sunfloweria Ramose
Asselian Hayasakapora akiyoshiensis Cylindrical stem
Akiyoshi Limestone Gp -
Triticites simplex zone Hayasakapora isaenis Cylindrical stem
Asselian Copacabana Fm Meekopora prosseri Bifurcating frond
Fistulipora cf. timorensis Encrusting
222
Age Formation Genus Species Morphology
Fistulipora incrustans Encrusting
Fistulipora sp. B Encrusting
Fistulipora sp. C Encrusting
Fistulipora titicacaensis Encrusting
Rhombopora corticata Erect
Rhombopora lepidodendroides Erect
Alternifenestella aff. pajerensis Fenestrate
Septopora andeana Fenestrate
Alternifenestella aspera Fenestrate
Minilya binodata Fenestrate
Alternifenestella cervoides Fenestrate
Polypora cf. megastoma Fenestrate
Polypora cyclopora Fenestrate
Polypora elliptica Fenestrate
Alternifenestella minor Fenestrate
Goniocladia peruviana Fenestrate
Fabifenestella sp. Fenestrate
Polypora sp. B Fenestrate
Tabulipora sp. Lamellate
Fistulipora carrascoi Ramose
Tabulipora cf. carbonaria Ramose
Streblotrypa sp. Ramose
Rhombotrypella typica Ramose
Asselian Copacabana Fm - Section 7 Goniocladia peruviana Fenestrate
Asselian Copacabana Fm - Section 8 Meekopora cf. prosseri Bifoliate frond
Polypora andina Fenestrate
Acanthocladia biserialis Fenestrate
Cervella cervoidea Fenestrate
223
Age Formation Genus Species Morphology
Fenestella huascatayana Fenestrate
Silvaseptopora incaica Fenestrate
Polypora inimica Fenestrate
Fenestella picchuensis Fenestrate
Polypora spissa Fenestrate
Asselian Copacabana Fm - Section 9 Meekopora cf. prosseri Bifoliate frond
Polypora andina Fenestrate
Cervella aspera Fenestrate
Silvaseptopora incaica Fenestrate
Fenestella pajerensis Fenestrate
Goniocladia peruviana Fenestrate
Polypora sp. Fenestrate
Polypora spissa Fenestrate
Asselian Elephant Canyon Fm Fistulipora sp. Cylindrical
Asselian Kapp Dunér Fm Rhombotrypella cf. stuckenbergi Ramose
Ascopora sterlitamakensis Ramose
Asselian Miharano Fm Polypora eliasi Fenestrate
Anastomopora orientalis Fenestrate
Minilya taishakuensis Fenestrate
Streblascopora lineata Ramose
Fistulipora miharanoensis Ramose
Asselian Nordenskiölbreen Fm -
Tyrrellfjellet Mbr
Eridopora sp. Encrusting
Alternifenestella bifida Fenestrate
Polypora kutorgae Fenestrate
224
Age Formation Genus Species Morphology
Rectifenestella veneris Fenestrate
Rectifenestella cf. ornata Fenestrate
Penniretepora cf. subtila Fenestrate
Coscinium cyclops Fenestrate
Flexifenestella grandis Fenestrate
Rectifenestella microporata Fenestrate
Ramipora minuta Fenestrate
Fabifenestella permiana Fenestrate
Polypora sp. Fenestrate
Archimedes sp. B Fenestrate
Polypora sulaensis Fenestrate
Tabulipora sp. Multilaminar,
Encrusting
Acanthocladia cf. rhombicellata Pinnate
Rhombotrypella dvinensis Ramose
Streblotrypa fasciculata Ramose
Ascopora grandis Ramose
Ascoporella grandis Ramose
Rhombotrypella invulgata Ramose
Rhombotrypella rectangulata Ramose
Rhabdomeson sp. Ramose
Timanodictya sp. A Ramose, Bifoliate
Ascopora sterlitamakensis Ramose
Ascopora muromensis Ramose
Asselian Omi Limestone Coscinotrypa minor Bifoliate
Tabulipora cf. maculosa Erect
Fistulipora sp. E Lamellate
Thamniscus? problematicus Ramose
Sulcoretepora sp. Ramose
225
Age Formation Genus Species Morphology
Asselian Panther Seep Fm Septopora cf. spinulosa Fenestrate
"Fenestella" sp. Fenestrate
Penniretepora sp. Pinnate
cf. Rhombopora sp. Ramose
Sakmarian unnamed formation - Diamond
Range
Timanodictya sp. Erect
Sakmarian Lyons Gp Polypora lyndoni Erect
Stenopora dickinsi Ramose
Stenopora fisheri Ramose
Stenopora lyndoni Ramose
Sakmarian Sunflower Fm Sulcoretepora corticata Bifoliate
Sulcoretepora irregularis Bifoliate frond
Rhabdomeson sp. Cylindrical
Fistulipora bifoliatus Erect bifoliate
Polypora sp. Fenestrate
Fistulipora lamellatis Lamellate
Penniretepora sp. Pinnate
Rhombopora cornwallis Ramose
Rhombotrypella sunfloweria Ramose
Sakmarian Abdullino Reef Hederella carbinarua Ramose
Sakmarian Akiyoshi Limestone Gp Fistulipora aff. grandis
volongensis
Lamellar
Fistulipora irimiensis Massive
Meekoporella akiyoshiensis Massive
226
Age Formation Genus Species Morphology
Stenopora toriyamai Massive
Streblascopora supergrossa Ramose
Sakmarian Copacabana Fm Meekopora sp. Bifoliate
Rhombopora kawabei Erect
Rhombopora lepidodendroides Erect
Polypora aff. inimica Fenestrate
Polypora cyclopora Fenestrate
Polypora elliptica Fenestrate
Polypora sp. A Fenestrate
Polypora sp. C Fenestrate
Septopora incaica Fenestrate
Septopora lineata Fenestrate
Dyscritella aff. komukensis Ramose
Dyscritella tenuirama Ramose
Fistulipora multidiaphragma Ramose
Pseudobatostom
ella
micropora Ramose
Pseudobatostom
ella
sp. Ramose
Pseudobatostom
ella
yanagidai Ramose
Rhombotrypella aff. gigantea Ramose
Rhombotrypella typica Ramose
Stenodiscus altiplana Ramose
Sakmarian Copacabana Fm - Section 7 Goniocladia peruviana Fenestrate
Sakmarian Copacabana Fm - Section 8 Meekopora cf. prosseri Bifoliate frond
Acanthocladia biserialis Fenestrate
227
Age Formation Genus Species Morphology
Cervella cervoidea Fenestrate
Fenestella huascatayana Fenestrate
Fenestella picchuensis Fenestrate
Polypora andina Fenestrate
Polypora inimica Fenestrate
Polypora spissa Fenestrate
Silvaseptopora incaica Fenestrate
Sakmarian Copacabana Fm - Section 9 Meekopora cf. prosseri Bifoliate frond
Cervella aspera Fenestrate
Fenestella pajerensis Fenestrate
Goniocladia peruviana Fenestrate
Polypora andina Fenestrate
Polypora sp. Fenestrate
Polypora spissa Fenestrate
Silvaseptopora incaica Fenestrate
Sakmarian Darlington Limestone Stenopora tasmaniensis Ramose
Sakmarian unnamed formation - Diamond
Range
Anisotrypella sp. Erect
Sakmarian Dwyka Fm Dyscritella cf. spinigera Cylindrical
Sakmarian Nelynya-shor Fm Paraseptopora uralica Pinnate
Anisotrypella radix Ramose
Uralotrypa lepida Ramose
Sakmarian Hinaclema svalbardensis Encrusting
Nordenskiölbreen Fm -
Tyrrellfjellet Mbr Alternifenestella bifida Fenestrate
228
Age Formation Genus Species Morphology
Polypora kutorgae Fenestrate
Rectifenestella veneris Fenestrate
Archimedes sp. B Fenestrate
Coscinium cyclops Fenestrate
Fabifenestella permiana Fenestrate
Flexifenestella grandis Fenestrate
Polypora martis Fenestrate
Ramipora minuta Fenestrate
Rectifenestella microporata Fenestrate
Ascopora grandis Ramose
Ascopora sp. Ramose
Ascopora sterlitamakensis Ramose
Ascoporella? sp. A Ramose
Rhombopora sp. A Ramose
Rhombotrypella invulgata Ramose
Sakmarian Robledo Mountains Fm Fistulipora sp. Encrusting
Tabulipora sp. Encrusting,
Ramose
Sakmarian Speiser Shale Fenestella spinulosa Fenestrate
Fenestella tenax Fenestrate
Minilya binodata Fenestrate
Protoretepora elliptica Fenestrate
Septopora spinulosa Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora auernigiana Pinnate
Penniretepora curvula Pinnate
Penniretepora flexistriata Pinnate
Rhombopora lepidodendroides Ramose
229
Age Formation Genus Species Morphology
Syringoclemis wrefordensis Ramose
Tabulipora carbonaria Ramose
Sakmarian Trogkofel Fm Eridopora ignota Encrusting
Eridopora sp. Encrusting
Stenophragmidi
um
lamellatum Encrusting
Carnocladia fascicularia Erect
Penniretepora trapezoida Erect
Prismopora sp. Erect
Alternifenestella subquadratopora Fenestrate
Alternifenestella tuberculifera Fenestrate
Paraptylopora sp. Pinnate
Penniretepora sp Pinnate
Primorella serena Ramose
Rhabdomeson hirtum Ramose
Rhombopora sp. Ramose
Streblascopora germana Ramose
Polypora sigillata Reticulated
Sakmarian Eridopora sp. Encrusting
Tabulipora sp. Encrusting
Prismopora digitata Erect
Carnocladia fascicularia Erect
Carnocladia punctata Erect
Alternifenestella aff. tenuiseptata Fenestrate
Alternifenestella nalivkini Fenestrate
Alternifenestella tenuiseptata Fenestrate
Upper Pseudoschwagerina
Limestone
Alternifenestella tuberculifera Fenestrate
Sakmarian Wasp Head Fm Stenopora spiculata Ramose
230
Age Formation Genus Species Morphology
Sakmarian Wreford Fm - Havensville
Limestone Mbr
Fistulipora carbonaria Encrusting sheets
Fistulipora incrustans Encrusting sheets
Fenestella spinulosa Fenestrate
Fenestella tenax Fenestrate
Minilya binodata Fenestrate
Septopora spinulosa Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora curvula Pinnate
Rhombopora lepidodendroides Ramose
Syringoclemis wrefordensis Ramose
Tabulipora carbonaria Ramose
Sakmarian Minilya binodata Fenestrate
Fistulipora carbonaria Encrusting sheets
Fistulipora incrustans Encrusting sheets
Meekopora prosseri Erect bifoliate
Fenestella tenax Fenestrate
Polypora aestacella Fenestrate
Polypora nodolinearis Fenestrate
Protoretepora elliptica Fenestrate
Septopora spinulosa Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora curvula Pinnate
Penniretepora flexistriata Pinnate
Rhombopora lepidodendroides Ramose
Syringoclemis wrefordensis Ramose
Wreford Fm - Schroyer
Limestone Mbr
Tabulipora carbonaria Ramose
231
Age Formation Genus Species Morphology
Sakmarian Fistulipora incrustans Encrusting sheets
Wreford Fm - Threemile
Limestone Mbr Fenestella spinulosa Fenestrate
Fenestella tenax Fenestrate
Minilya binodata Fenestrate
Protoretepora elliptica Fenestrate
Septopora spinulosa Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora curvula Pinnate
Filiramoporina kretaphilia Ramose
Rhombopora lepidodendroides Ramose
Syringoclemis wrefordensis Ramose
Artinskian Aiduna Fm - Location A Hyphasmopora katoi Cylindrical
Saffordotaxis cf. wanneri Cylindrical
Streblotrypa elegans Cylindrical
Polypora sp. Fenestrate
Polypora timorensis Fenestrate
Fenestella" spp. Fenestrate
Acanthocladia cf. regularis Pinnate
Ascopora nakornsrii Ramose
Rhabdomeson mammillatun Ramose
Streblascopora cf. fasciculata Ramose
Streblascopora irianica Ramose
Fistulipora cf. timorensis Erect
Goniocladia timorensis Fenestrate
Artinskian Aiduna Fm - Location B Clausotrypa cf. conferta Cylindrical
Rhabdomeson cf. grande Ramose
Eridopora parasitica Encrusting
Dyscritella adnascens Lamellar
232
Age Formation Genus Species Morphology
Fistulipora labratula Massive
Fistulipora lunatifera Massive
Artinskian Akiyoshi Limestone Gp -
Pseudofusulina ambigua zone
Fistulipora aff. grandis
volongensis
Lamellar
Fistulipora irimiensis Massive
Meekoporella akiyoshiensis Massive
Stenopora toriyamai Massive
Streblascopora supergrossa Ramose
Artinskian Stenopora ovata Dendroid,
Encrusting
Levifenestella expansa Fenestrate
Parapolypora sp. A Fenestrate
Polypora virga Fenestrate
Rectifenestella granulifera Fenestrate
Stenopora berriedalensis Ramose
Stenopora crinita Ramose
Berriedale Limestone
Stenopora etheridgei Ramose
Artinskian Buffel Fm Dyscritella eximia Dendroid
Dyscritella trivialis Dendroid
Dyscritellina tecta Encrusting
Eridopora rara Encrusting
Metelipora improvisa Encrusting
Alternifenestella horologia Fenestrate
Goniocladia immensa Fenestrate
Rectifenestella bowenensis Fenestrate
Diploporaria sp. Pinnate
Crockfordia multinodata Ramose
233
Age Formation Genus Species Morphology
Stenopora ovata Ramose
Stenopora tasmaniensis Ramose
Streblascopora sp. Ramose
Paramaychella spinosa Ramose
Artinskian Fistulipora? sp. Encrusting,
Massive
Timanodictya? sp. Erect bifoliate
Polypora aff. bassleri Fenestrate
Polypora aff. halliana Fenestrate
Polypora aff. keyserlingi Fenestrate
Polypora aff. sparsa Fenestrate
Polypora cf. daurica Fenestrate
Cathedral Mountain Fm -
Poplar Tank Mbr
Polypora cf. novella Fenestrate
Polypora cf. subborealis Fenestrate
Polypora cf. virga Fenestrate
Polypora hirsuta Fenestrate
Polypora mexicana Fenestrate
Polypora nodocarinata Fenestrate
Polypora perturbata Fenestrate
Polypora robustoformis Fenestrate
Polypora sp. Fenestrate
Polypora sp. 1 Fenestrate
Polyporella cf. ulakhanensis Fenestrate
Protoretepora aff. elliptica Fenestrate
Protoretepora aff. fujimotoi Fenestrate
Protoretepora aff. vitiosa Fenestrate
Protoretepora cf. ampla Fenestrate
Protoretepora cf. crassa Fenestrate
Protoretepora cf. praepluriformis Fenestrate
234
Age Formation Genus Species Morphology
Protoretepora soyanensis Fenestrate
Protoretepora sp. Fenestrate
Septopora aff. phyllata Fenestrate
Septopora aff. pinnata Fenestrate
Septopora cf. flabellata Fenestrate
Septopora orientalis Fenestrate
Septopora sp. Fenestrate
Synocladia sp. 1 Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora sp. Pinnate
Rhadomeson? sp. Ramose
Rhombopora lepidodendroides Ramose
Rhombopora sp. Ramose
Streblascopora sp. Ramose
Tabulipora sp. Ramose
Ulrichotrypa? sp. Ramose
Artinskian Cattle Creek Fm Fenestella affluensa Fenestrate
Fenestella cf. columnaris Fenestrate
Fenestella cf. dispersa Fenestrate
Fenestella dispersa Fenestrate
Fenestella fossula Fenestrate
Fenestella horologia Fenestrate
Fenestella simulatrix Fenestrate
Polypora pertinax Fenestrate
Polyporella woodsi Fenestrate
Protoretepora ampla Fenestrate
Diploporaria sp. Pinnate?
Artinskian Chuping Limestone - Bukit Fenestella cf. retiformis Fenestrate
235
Age Formation Genus Species Morphology
Mata Ayer
Artinskian Chuping Limestone – Pulau
Jonk
Cyclotrypa alexanderi Cylindrical
Fistulipora hupehensis Cylindrical
Polypora cf. timorensis Fenestrate
Polypora gigantea Fenestrate
Counsel Creek Fm Levifenestella expansa Fenestrate
Rectifenestella counselensis Fenestrate
Stenopora grantonensis Ramose
Artinskian Deep Bay Fm Polypora rebarbensis Fenestrate
Shulgapora magnafenestrata Fenestrate
Artinskian Dwyka Fm Dyscritella cf. spinigera Cylindrical
Artinskian Gipshuken Fm Hinaclema svalbardensis Encrusting
Polypora martis Fenestrate
Polypora voluminosa Fenestrate
Artinskian Great Bear Cape Fm Stenophragmidi
um
sp. Encrusting
Acanthocladia cf. rhombicellata Fenestrate
Acanthocladia cf. sparsifurcata Fenestrate
Alternifenestella bifida Fenestrate
Alternifenestella cf. invisitata Fenestrate
Alternifenestella crassiseptata Fenestrate
Alternifenestella cyclotriangulata Fenestrate
Bashkirella operculata Fenestrate
236
Age Formation Genus Species Morphology
Fabifenestella cf. subvirgosa Fenestrate
Fabifenestella cf. virgosa Fenestrate
Fabifenestella tortuosa Fenestrate
Fenestella akselensis Fenestrate
Penniretepora invisa Fenestrate
Polypora confirmata Fenestrate
Polypora kossjensis Fenestrate
Polypora kutorgae Fenestrate
Polypora martis Fenestrate
Polypora voluminosa Fenestrate
Rectifenestella microporata Fenestrate
Rectifenestella robusta Fenestrate
Primorella superba Ramose
Primorella tundrica Ramose
Streblascopora germana Ramose
Streblascopora vera Ramose
Artinskian Hambergfjellet Fm Dyscritella cf. narjanmarica Ramose
Streblascopora germana Ramose
Timanodictya dichotoma Ramose
Artinskian Dyscritella sp. A Encrusting
Girtyporina sp. Encrusting
Meekopora magnusi Erect bifoliate
Alternifenestella bifida Fenestrate
Alternifenestella cf. greenharbourensis Fenestrate
Alternifenestella cf. minuscula Fenestrate
Kapp Starostin Fm - Vøringen
Mbr 5y
Alternifenestella sp. A Fenestrate
Alternifenestella subquadratopora Fenestrate
Fabifenestella sp. A Fenestrate
237
Age Formation Genus Species Morphology
Fenestella akselensis Fenestrate
Fenestella reversicnotta Fenestrate
Goniocladia sp. Fenestrate
Lyropora serissima Fenestrate
Polypora brevicellata Fenestrate
Polypora martis Fenestrate
Polyporella biarmica Fenestrate
Polyporella borealis Fenestrate
Polyporella sp. Fenestrate
Polyporella subcrotilla Fenestrate
Ramipora cf. hochstetteri Fenestrate
Rectifenestella araxensis Fenestrate
Rectifenestella microporata Fenestrate
Rectifenestella retiformis Fenestrate
Rectifenestella sp. A Fenestrate
Rectifenestella sp. B Fenestrate
Clausotrypa monticola Ramose
Cyclotrypa distincta Ramose,
Encrusting
Cyclotrypa eximina Ramose,
Encrusting
Dyscritella bogatensis Ramose
Dyscritella minuta Ramose
Dyscritella narjanmarica Ramose
Permoheloclema merum Ramose
Primorella cf. polita Ramose
Rhombotrypella alfredensis Ramose
Rhombotrypella arbuscula Ramose
Stenopora thula Ramose
Streblascopora germana Ramose
238
Age Formation Genus Species Morphology
Tabulipora greenlandensis Ramose
Tabulipora siedleckii Ramose
Tabulipora sp. A Ramose
Timanodictya nikifororvae Ramose
Artinskian Kim Fjelde Fm Tabulipora sp. Ramose
Artinskian Nabeyama Fm Rhombopora? sp. Erect
Stenopora sp. A Erect
Fistulipora regularis Lamellate
Prismopora kuzuensis Triradiate?
Artinskian Poole Sandstone - Nura Nura
Mbr
Streblotrypa marmionensis Ramose
Artinskian Dyscritella sp. Cylindrical
Dyscritella sp. Cylindrical
Fenestella basleoensis Fenestrate
Fenestella cf. basleoensis Fenestrate
Fenestella cf. procera Fenestrate
Fenestella chapmani Fenestrate
Fenestella horologia Fenestrate
Fenestella krachokensis Fenestrate
Fenestella sp. A Fenestrate
Fenestella sp. B Fenestrate
Polypora quadricella Fenestrate
Rat Buri Ls - Khao Chong
Krachok
Thamniscus sp. Fenestrate
Artinskian Rat Buri Ls - Khao Phrik Hexagonella khaophrikensis Bifoliate
Hexagonella robusta Bifoliate
239
Age Formation Genus Species Morphology
Streblascopora ratburiensis Cylindrical
Streblascopora sp. Cylindrical
Streblotrypa? crassa Cylindrical
Acanthocladia thaiensis Fenestrate
Fenestella megacapillaris Fenestrate
Polypora aff. variocellata Fenestrate
Polypora sp. Fenestrate
Hexagonella kobayashii Frondose
Ascopora asiatica Ramose
Ascopora magna Ramose
Ascopora nakornsrii Ramose
Ascopora sp. Ramose
Ascopora yanagidae Ramose
Fistulipora horowitzi Ramose
Artinskian Rat Buri Ls - Ko Muk Dyscritella cf. tenuirama Cylindrical
Dyscritella komukensis Cylindrical
Dyscritella sp. A Cylindrical
Dyscritella tchurkensis Cylindrical
Streblascopora exillis Cylindrical
Sulcoretepora sp. Erect
Sulcoretepora thailandica Erect
Fenestella cf. horologia Fenestrate
Fenestella jabiensis Fenestrate
Fenestella komalarjuni Fenestrate
Fenestella megacapillaris Fenestrate
Fenestella sp. Fenestrate
Fenestella thaiensis Fenestrate
Fenestella
(Minilya)
pseudoamplia Fenestrate
240
Age Formation Genus Species Morphology
Goniocladia timorensis Fenestrate
Polypora fritzi Fenestrate
Polypora multiporiferata Fenestrate
Fistulipora lamella Hemispherical
Fistulipora satoi Massive,
Encrusting
Fistulipora simillina Massive
Penniretepora pecularis Pinnate
Penniretepora siamensis Pinnate
Penniretepora tropica Pinnate
Ascopora asiatica Ramose
Fistulipora hamadae Ramose
Fistulipora horowitzi Ramose
Fistulipora komukensis Ramose
Fistulipora sp. Ramose
Fistulipora tenella Ramose
Goniocladia laxa Ramose
Rhabdomeson mammilatum Ramose
Streblascopora cf. marmionensis Ramose
Streblascopora komukensis Ramose
Streblotrypa elegans Ramose
Streblotrypa? thaiensis Ramose
Artinskian Sarga Fm Synocladia georgii Fenestrate
Artinskian Trogkofel Fm Eridopora ignota Encrusting
Eridopora sp. Encrusting
Stenophragmidi
um
lamellatum Encrusting
Prismopora sp. Erect
241
Age Formation Genus Species Morphology
Carnocladia fascicularia Erect
Penniretepora trapezoida Erect
Alternifenestella subquadratopora Fenestrate
Alternifenestella tuberculifera Fenestrate
Paraptylopora sp. Pinnate
Penniretepora sp Pinnate
Primorella serena Ramose
Rhabdomeson hirtum Ramose
Rhombopora sp. Ramose
Streblascopora germana Ramose
Polypora sigillata reticulated
Kungurian Meekoporella akiyoshiensis Bifoliate
Pseudobatostom
ella
sp. Cylindrical
Hayasakapora sp. Cylindrical stem
Rhabdomeson ofukuensis Cylindrical stem
Fistulipora ozawae Encrusting
Leioclema micropora Encrusting
Leioclema muratae Encrusting
Fenestella macronodata Fenestrate
Fenestella otae Fenestrate
Fenestella
(Loculiporina)
toriyamae Fenestrate
Polypora ovalifenestrata Fenestrate
Penniretepora hashimotoi Pinnate
Streblascopora diaphragma Ramose
Akiyoshi Limestone Gp -
Parafusulina kaerimizensis
zone
Streblascopora supergrossa Ramose
242
Age Formation Genus Species Morphology
Kungurian Balikelik (Baliqliq) Fm Ascopora delicata Cylindrical
Eridopora? uncata Cylindrical
Rhabdomeson consimile Cylindrical
Rhabdomeson cylindricum Cylindrical
Rhabdomeson grande Cylindrical
Rhabdomeson mammillata Cylindrical
Streblotrypa grandis Cylindrical
Tabulipora xinjiangensis Cylindrical,
Massive
Dybowskiella sp. Encrusting
Fistulipora gigantea lemellaris Encrusting
Fenestella cf. petschorica Fenestrate
Fenestella kalpingensis Fenestrate
Fenestella kungurensis Fenestrate
Fenestella nikiforovae Fenestrate
Fenestella retiformis Fenestrate
Fenestella sp. Fenestrate
Synocladia robusta Fenestrate
Fistulipora gigantea Massive
Penniretepora granulata Pinnate
Dybowskiella hupehensis Ramose
Fistulipora confluens Ramose
Klaucena elliptica Ramose, Bifoliate
Maychella rhomboidea Ramose
Rhabdomeson commune Ramose
Rhabdomeson mammillata ellipticus Ramose
Rhabdomeson xinjiangensis Ramose
Streblascopora abnorme Ramose
Streblascopora cf. marmionensis Ramose
Streblascopora fasciculata Ramose
243
Age Formation Genus Species Morphology
Streblotrypa sinensis Ramose
Fistulipora insidians Subramose
Kungurian Cathedral Mountain Fm Fistulipora? sp. Encrusting,
Massive
Timanodictya? sp. Erect bifoliate
Fenestella amplia Fenestrate
Fenestella cf. schucherti Fenestrate
Fenestella firmior Fenestrate
Fenestella girtyi Fenestrate
Fenestella spinulifera Fenestrate
Polypora aff. bassleri Fenestrate
Polypora andina Fenestrate
Polypora cf. novella Fenestrate
Polypora sp. Fenestrate
Protoretepora aff. elliptica Fenestrate
Protoretepora cf. praepluriformis Fenestrate
Septopora aff. robusta Fenestrate
Septopora cf. biserialis Fenestrate
Septopora orientalis Fenestrate
Septopora sp. 1 Fenestrate
Septopora sp. 4 Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora sp. Pinnate
Rhabdomeson? sp. Ramose
Rhombopora lepidodendroides Ramose
Syringoclemis? sp. Ramose
Tabulipora carbonaria Ramose
244
Age Formation Genus Species Morphology
Kungurian Fistulipora? sp. Encrusting,
Massive
Timanodictya? sp. Erect bifoliate
Fenestella girtyi Fenestrate
Cathedral Mountain Fm -
Sullivan Peak Mbr
Acanthocladia guadalupensis Pinnate
Penniretepora sp. Pinnate
Ulrichotrypa? sp. Ramose
Kungurian Timanodictya? sp. Erect bifoliate
Anastomopora cf. orientalis Fenestrate
Fenestella hilli Fenestrate
Fenestella spinulifera Fenestrate
Polypora andina Fenestrate
Polypora robustoformis Fenestrate
Polypora sp. Fenestrate
Septopora aff. robusta Fenestrate
Septopora aff. spinulosa Fenestrate
Septopora cf. biserialis Fenestrate
Septopora orientalis Fenestrate
Septopora sp. Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora sp. Pinnate
Cathedral Mountain Fm -
Wedin Conglomerate Mbr
Rhombopora? sp. Ramose
Kungurian Cyclotrypa alexanderi Cylindrical
Fistulipora hupehensis Cylindrical
Polypora cf. timorensis Fenestrate
Chuping Limestone – Pulau
Jonk
Polypora gigantea Fenestrate
245
Age Formation Genus Species Morphology
Kungurian Dzhiakun’skaya Fm Hayasakapora ambigua Ramose
Hayasakapora erectoradiata Ramose
Timanodictya ellipsoidalis Ramose
Kungurian Hambergfjellet Fm Streblascopora germana Ramose
Timanodictya dichotoma Ramose
Kungurian Kapp Starostin Fm - Vøringen
Mbr
Dyscritella sp. A Encrusting
Girtyporina sp. Encrusting
Meekopora magnusi erect bifoliate
Alternifenestella bifida Fenestrate
Alternifenestella cf. greenharbourensis Fenestrate
Alternifenestella cf. minuscula Fenestrate
Alternifenestella sp. A Fenestrate
Alternifenestella subquadratopora Fenestrate
Fabifenestella sp. A Fenestrate
Fenestella akselensis Fenestrate
Fenestella reversicnotta Fenestrate
Goniocladia sp. Fenestrate
Lyropora serissima Fenestrate
Polypora brevicellata Fenestrate
Polypora martis Fenestrate
Polyporella biarmica Fenestrate
Polyporella borealis Fenestrate
Polyporella sp. Fenestrate
Polyporella subcrotilla Fenestrate
Ramipora cf. hochstetteri Fenestrate
Rectifenestella araxensis Fenestrate
Rectifenestella microporata Fenestrate
246
Age Formation Genus Species Morphology
Rectifenestella retiformis Fenestrate
Rectifenestella sp. A Fenestrate
Rectifenestella sp. B Fenestrate
Clausotrypa monticola Ramose
Cyclotrypa distincta Ramose,
Encrusting
Cyclotrypa eximina Ramose,
Encrusting
Dyscritella bogatensis Ramose
Dyscritella minuta Ramose
Dyscritella narjanmarica Ramose
Permoheloclema merum Ramose
Primorella cf. polita Ramose
Rhombotrypella alfredensis Ramose
Rhombotrypella arbuscula Ramose
Stenopora thula Ramose
Streblascopora germana Ramose
Tabulipora greenlandensis Ramose
Tabulipora siedleckii Ramose
Tabulipora sp. A Ramose
Timanodictya nikifororvae Ramose
Kungurian Kim Fjelde Fm Tabulipora sp. Ramose
Kungurian Kozhim Fm Cyclotrypa aperta Encrusting?
Dyscritella epidema Ramose
Neoeridotrypella astrica Ramose
Kungurian Loray Fm Fenestella sp. Fenestrate
Tabulipora sp. Ramose
247
Age Formation Genus Species Morphology
Kungurian Miseryfjellet Fm Fenestella paratuberculifera Fenestrate
Ramipora hochstetteri Fenestrate
Dyscritella bogatensis Ramose
Gilmouropora heintzi Ramose
Rhombotrypella alfredensis Ramose
Stenopora dickinsi Ramose
Stenopora jungersenensis Ramose
Timanodictya nikifororvae Ramose
Tabulipora greenlandensis Ramose
Kungurian Nooncanbah Fm Fistulamina lata Erect, Bifoliate
Streblotrypa marmionensis Ramose
Australopolypora fovea Fenestrate
Fenestella cacuminatis Fenestrate
Fenestella columnaris Fenestrate
Fenestella disjecta Fenestrate
Fenestella horologia Fenestrate
Fenestella lennardi Fenestrate
Fenestella rudiacarinata Fenestrate
Fenestella valentis Fenestrate
Minilya duplaris Fenestrate
Minilya princeps Fenestrate
Polypora multiporifera Fenestrate
Polypora sp. Fenestrate
Kungurian Oxtrack Fm Alternifenestella spinifera Fenestrate
Polyporella magnifenestrata Fenestrate
Rectifenestella expansa Fenestrate
Rectifenestella granulifera Fenestrate
Penniretepora triporosa Pinnate
248
Age Formation Genus Species Morphology
Crockfordia multinodata Ramose/Massive
Stenopora ovata Ramose/Encrusting
Stenopora tasmaniensis Ramose
Kungurian Road Canyon Fm Fistulipora? sp. Encrusting
Timanodictya? sp. Erect bifoliate
Anastomopora cf. orientalis Fenestrate
Fenestella aff. spinulosa Fenestrate
Fenestella archimediformis Fenestrate
Fenestella firmior Fenestrate
Fenestella hilli Fenestrate
Fenestella sp. Fenestrate
Isotrypa sp. Fenestrate
Parafenestralia cf. gregalis Fenestrate
Polypora aff. hivatchensis Fenestrate
Polypora aff. sparsa Fenestrate
Polypora balkhaschensiformis Fenestrate
Polypora cf. darashamensis Fenestrate
Polypora cf. daurica Fenestrate
Polypora cf. hinganensis Fenestrate
Polypora cf. irregularis Fenestrate
Polypora hirsuta Fenestrate
Polypora mexicana Fenestrate
Polypora perturbata Fenestrate
Polypora robustoformis Fenestrate
Polypora sp. Fenestrate
Polypora sp. 1 Fenestrate
Polypora sp. 2 Fenestrate
Protoretepora aff. elliptica Fenestrate
Protoretepora aff. vitiosa Fenestrate
249
Age Formation Genus Species Morphology
Protoretepora cf. ampla Fenestrate
Protoretepora soyanensis Fenestrate
Protoretepora sp. Fenestrate
Septopora aff. lineata Fenestrate
Septopora aff. pinnata Fenestrate
Septopora aff. robusta Fenestrate
Septopora aff. spinulosa Fenestrate
Septopora cf. biserialis Fenestrate
Septopora cf. blanda Fenestrate
Septopora cf. cestriensis Fenestrate
Septopora cf. flabellata Fenestrate
Septopora cf. quasiorientalis Fenestrate
Septopora orientalis Fenestrate
Septopora sp. 1 Fenestrate
Septopora sp. 10 Fenestrate
Septopora sp. 11 Fenestrate
Septopora sp. 2 Fenestrate
Septopora sp. 3 Fenestrate
Septopora sp. 5 Fenestrate
Septopora sp. 6 Fenestrate
Septopora sp. 7 Fenestrate
Septopora sp. 8 Fenestrate
Septopora sp. 9 Fenestrate
Synocladia sp. 1 Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora sp. Pinnate
Acanthoclema sp. Ramose
Rhabdomeson bellum Ramose
Rhombopora? sp. Ramose
Stenodiscus? sp. Ramose
250
Age Formation Genus Species Morphology
Stenopora? sp. Ramose
Tabulipora sp. Ramose
Ulrichotrypa? sp. Ramose
Kungurian Shoushangou Fm Hayasakapora sp. Ramose
Kungurian Toroweap Fm Dyscritella muddyensis Dendroid
Hexagonella cf. recta Dendroid
Rhabdomeson petita Dendroid
Stenodiscus sp. Dendroid
Stenopora torowensis Dendroid
Timanodictya cf. dichotoma Dendroid
Penniretepora oppositus Fenestrate
Polypora rhomboidensis Fenestrate
Polypora sargaensis Fenestrate
Reteporidra anaphora Fenestrate
Septopora bilateralis Fenestrate
Wjatkella permiana Fenestrate
Meekopora sp. Lamellar
Cyclotrypa cf. multiformis Massive
Fistulipora cf. enodata Massive
Streblotrypa regularis Ramose
Kungurian Wandrawandian Siltstone Fenestella sp. A Fenestrate
Laxifenestella exserta Fenestrate
Laxifenestella oviferosa Fenestrate
Levifenestella altacarinata Fenestrate
Minilya bituberculata Fenestrate
Paucipora ulladullaensis Fenestrate
Polypora dichotoma? Fenestrate
251
Age Formation Genus Species Morphology
Rectifenestella sparsa Fenestrate
Shulgapora magnafenestrata Fenestrate
Dyscritella espinensis Ramose
Stenopora seriatensis Ramose
Kungurian Yongde Fm Nikiforovella gigantea Ramose
Kungurian Zhuravlev strata Dyscritella cf. lucida Ramose
Roadian Balikelik (Baliqliq) Fm Ascopora delicata Cylindrical
Eridopora? uncata Cylindrical
Rhabdomeson consimile Cylindrical
Rhabdomeson cylindricum Cylindrical
Rhabdomeson grande Cylindrical
Rhabdomeson mammillata Cylindrical
Streblotrypa grandis Cylindrical
Tabulipora xinjiangensis Cylindrical,
Massive
Dybowskiella sp. Encrusting
Fistulipora gigantea lemellaris Encrusting
Fenestella kalpingensis Fenestrate
Fenestella kungurensis Fenestrate
Fenestella nikiforovae Fenestrate
Fenestella retiformis Fenestrate
Fenestella cf. petschorica Fenestrate
Fenestella sp. Fenestrate
Synocladia robusta Fenestrate
Fistulipora timorensis Lamellate
Fistulipora gigantea Massive
Dybowskiella hupehensis Ramose
252
Age Formation Genus Species Morphology
Penniretepora granulata Pinnate
Fistulipora confluens Ramose
Rhabdomeson commune Ramose
Rhabdomeson mammillata ellipticus Ramose
Maychella rhomboidea Ramose
Rhabdomeson xinjiangensis Ramose
Streblascopora abnorme Ramose
Streblascopora cf. marmionensis Ramose
Streblascopora fasciculata Ramose
Streblotrypa sinensis Ramose
Klaucena elliptica Ramose, Bifoliate
Fistulipora insidians Subramose
Roadian Coolkilya Sandstone Fenestella disjecta Fenestrate
Roadian Iwaizaki Fm - Unit 4 Meekopora delicata Bifoliate
Sulcoretepora nipponica Bifoliate
Fistulipora cf. timorensis Encrusting
Hayasakapora erectoradiata Erect cylindrical
Pseudobatostom
ella
spinigera Erect cylindrical
Saffordotaxis morikawae Erect cylindrical
Fenestella rhomboidea Fenestrate
Fistulipora sp. C Lamellate
Streblascopora delicatula Ramose
Roadian Kaibab Fm - Beta Mbr Meekopora parilis Bifoliate
Streblotrypa sp. Dendroid
Bicorbis arizonica Fenestrate
?Stenodiscus sp. Ramose
253
Age Formation Genus Species Morphology
Girtypora maculata Ramose
Rhabdomeson sp. Ramose
Roadian Kapp Starostin Fm Timanodictya nikifororvae Erect bifoliate,
Ramose
Ramipora hochstetteri Fenestrate
Neoeridotrypella sp. Multilamellar
Stenopora timanensis Multilamellar
Anisotrypella sp. Multilamellar
Dyscritella arctica Multilamellar
Dyscritella savinaensis Multilamellar
Rhombotrypella alfredensis Multilamellar
Rhombotrypella insolita Multilamellar
Tabulipora aberrans Multilamellar
Tabulipora arcticensis Multilamellar
Stenopora jurgersenensis Ramose
Dyscritella spinigera Ramose
Primorella polita Ramose
Dyscritella parallela Ramose
Gilmouropora heintzi Ramose
Roadian Kozhim Rudnik Fm Iraidina ramosa Encrusting,
Bifoliate
Roadian Lightjack Fm Fenestella horologia Fenestrate
Streblascopora marmionensis Ramose
Roadian Minnie Point Fm Polypora rebarbensis Fenestrate
Roadian Road Canyon Fm Fistulipora? sp. Encrusting,
254
Age Formation Genus Species Morphology
Massive
Timanodictya? sp. Erect bifoliate
Anastomopora cf. orientalis Fenestrate
Fenestella aff. spinulosa Fenestrate
Fenestella archimediformis Fenestrate
Fenestella firmior Fenestrate
Fenestella hilli Fenestrate
Fenestella sp. Fenestrate
Isotrypa sp. Fenestrate
Parafenestralia cf. gregalis Fenestrate
Polypora aff. hivatchensis Fenestrate
Polypora aff. sparsa Fenestrate
Polypora balkhaschensiformis Fenestrate
Polypora cf. darashamensis Fenestrate
Polypora cf. daurica Fenestrate
Polypora cf. hinganensis Fenestrate
Polypora cf. irregularis Fenestrate
Polypora hirsuta Fenestrate
Polypora mexicana Fenestrate
Polypora perturbata Fenestrate
Polypora robustoformis Fenestrate
Polypora sp. Fenestrate
Polypora sp. 1 Fenestrate
Polypora sp. 2 Fenestrate
Protoretepora aff. elliptica Fenestrate
Protoretepora aff. vitiosa Fenestrate
Protoretepora cf. ampla Fenestrate
Protoretepora soyanensis Fenestrate
Protoretepora sp. Fenestrate
Septopora aff. lineata Fenestrate
255
Age Formation Genus Species Morphology
Septopora aff. pinnata Fenestrate
Septopora aff. robusta Fenestrate
Septopora aff. spinulosa Fenestrate
Septopora cf. biserialis Fenestrate
Septopora cf. blanda Fenestrate
Septopora cf. cestriensis Fenestrate
Septopora cf. flabellata Fenestrate
Septopora cf. quasiorientalis Fenestrate
Septopora orientalis Fenestrate
Septopora sp. 1 Fenestrate
Septopora sp. 10 Fenestrate
Septopora sp. 11 Fenestrate
Septopora sp. 2 Fenestrate
Septopora sp. 3 Fenestrate
Septopora sp. 5 Fenestrate
Septopora sp. 6 Fenestrate
Septopora sp. 7 Fenestrate
Septopora sp. 8 Fenestrate
Septopora sp. 9 Fenestrate
Synocladia sp. 1 Fenestrate
Acanthocladia guadalupensis Pinnate
Penniretepora sp. Pinnate
Acanthoclema sp. Ramose
Rhabdomeson bellum Ramose
Rhombopora? sp. Ramose
Stenodiscus? sp. Ramose
Stenopora? sp. Ramose
Tabulipora sp. Ramose
Ulrichotrypa? sp. Ramose
256
Age Formation Genus Species Morphology
Roadian Vladivostok Fm Iraidina damperovi Encrusting
Hayasakapora sp. Ramose
Ruzhencevia nevolinae Ramose
Fenestella columnaris Fenestrate
Fenestella disjecta Fenestrate
Fenestella horologia Fenestrate
Minilya duplaris Fenestrate
Australopolypora fovea Fenestrate
Polypora multiporifera Fenestrate
Polypora retificis Fenestrate
Wordian Iwaizaki Fm - Unit 6 Goniocladia intricata Bifoliate
Meekopora delicata Bifoliate
Meekopora densa Bifoliate
Fistulipora cf. timorensis Encrusting
Fistulipora iwaizakiensis Encrusting
Pseudobatostom
ella
spinigera Erect
Pseudobatostom
ella
igoi Ramose
Pseudobatostom
ella
yamazakii Ramose
Ramipora ambigua Ramose
Wordian Kanokura Fm Meekopora delicata Bifoliate
Sulcoretepora nipponica Ramose, Bifoliate
Fistulipora kesenumensis Encrusting
Fistulipora megastoma Encrusting
Leioclema globosa Encrusting
Thamniscus cf. dubius Ramose
257
Age Formation Genus Species Morphology
Fenestella cf. retiformis Fenestrate
Polypora endoi Fenestrate
Polypora fujimotoi Fenestrate
Polypora hataii Fenestrate
Polypora sugiyamae Fenestrate
Polypora toyokoae Fenestrate
Septopora kawamatae Fenestrate
Pseudobatostom
ella
sp. Ramose
Wordian Kapp Starostin Fm Timanodictya nikiforovae Ramose, Bifoliate
Ramipora hochstetteri Fenestrate
Neoeridotrypella sp. Multilamellar
Stenopora timanensis Multilamellar
Anisotrypella sp. Multilamellar
Dyscritella arctica Multilamellar
Dyscritella savinaensis Multilamellar
Rhombotrypella alfredensis Multilamellar
Rhombotrypella insolita Multilamellar
Tabulipora aberrans Multilamellar
Tabulipora arcticensis Multilamellar
Stenopora jurgersenensis Ramose
Dyscritella spinigera Ramose
Primorella polita Ramose
Dyscritella parallela Ramose
Gilmouropora heintzi Ramose
Wordian Malbina Fm Polypora rebarbensis Fenestrate
Stenopora berriedalensis Ramose
Stenopora crinita Ramose
258
Age Formation Genus Species Morphology
Wordian Maokou Fm Girtypora sp. Ramose
Hayasakapora sp. Ramose
Timanodictya zunyiensis Ramose
Wordian Mission Argillite Fm Dyscritella iwaizakiensis Dendroid
Fistuliramus pacificus Dendroid
Hayasakapora cf. erectoradiata Dendroid
Pamirella oculus Dendroid
Pinegopora petita Dendroid
Rhombotrypella kettlensis Dendroid
Sakagamiina easternensis Dendroid
Tabulipora colvillensis Dendroid
Coeloclemis urhausenii Encrusting
Neoeridotrypella missionensis Encrusting
Alternifenestella vagrantia Fenestrate
Mackinneyella stylettia Fenestrate
Polypora arbusca Fenestrate
Wjatkella nanea Fenestrate
Meekoporella inflecta Ramose
Wordian Murdock Mountain Fm Dyscritella acanthostylia Dendroid
Hinganella felderi Dendroid
Morozoviella praecurriensis Dendroid
Neoeridotrypella schilti Dendroid
Pseudobatostom
ella
irregularis Dendroid
Stenopora parvaexozona Dendroid, Encrusting
Streblotrypa elongata Dendroid
Kingopora cf. ehrenbergi Fenestrate
259
Age Formation Genus Species Morphology
Stenodiscus murdockensis Massive, Dendroid
Wordian Neoschwagerina Beds Dybowskiella elegantula Encrusting
Eridopora cf. parasitica Encrusting
Archimedes sp. Frondose
Stenopora obesa Massive, Encrusting
Dyscritella sp. Ramose
Streblascopora cf. diaphragma Ramose
Actinotrypella sp. Ramose bifoliate
Sulcoretepora? sp. Ramose bifoliate
Wordian Pietra di Salomone Fistulipora regularis Massive
Hayasakapora sp. Ramose
Wordian Puchenpra Fm Streblascopora cf. marmionensis Ramose
Wordian Ruteh Fm Rhombopora polyporata Cylindrical
Fenestella perelegans Fenestrate
Polypora koninckiana Fenestrate
Septopora sp. Fenestrate
Wordian unknown Fm - Kulaya River Girtyporina applicata Pinnate
Wordian unknown Fm - Volgograd Belt Ruzhencevia incrustata Ramose
Wordian unknown Fm - Vyatka River Girtypora ramosa Ramose
Wordian unknown Fm - Western
Kazakhstan
Ruzhencevia incrustata Ramose
260
Age Formation Genus Species Morphology
Wordian Vladivostok Fm Hayasakapora sp. Ramose
Ruzhencevia nevolinae Ramose
Wordian Xiangbo reef Dybowskiella sp. Encrusting
Fistulipora sp. Encrusting
Fenestella donaica Fenestrate
Fenestella sp. Fenestrate
Polypora guangxiensis Fenestrate
Polypora sp. Fenestrate
Ascopora cf. quadritubulata Ramose
Eridopora sp Ramose, Encrusting
Rhabdomeson sp. Ramose
Capitanian Abadeh Fm - Unit 4 Eridopora cf. parasitica Encrusting
Polypora abadehensis Encrusting
Polypora aff. darashamensis Encrusting
Polypora magnicava Encrusting
Polypora soyanensis Encrusting
Polypora striata Encrusting
Polypora tubulosa Encrusting
Septopora lineata Encrusting
Capitanian Hayasakapora yanagidai Cylindrical
Fistulipora octonaria Cylindrical
Rhabdomeson sp. Cylindrical
Eridopora cf. luminosa Encrusting
Fistulipora aff. lunatifera Encrusting
Fistulipora cf. megastoma Encrusting
Akiyoshi Limestone Gp -
Lepidolina multiseptata
shiraiensis zone
Paralioclema eplicatum Encrusting
261
Age Formation Genus Species Morphology
Fistulipora pseudolunaris Encrusting?
Girtyporina cf. applicata Foliate, Unilaminar
Hexagonella sp. Frondose
Clausotrypa monstruosa Massive, Cylindrical
Girtyporina crassa Massive, Cylindrical
Girtyporina gorjunovae Massive, Cylindrical
Permolioclema akiyoshiensis Massive, Encrusting
Dyscritella shigeyasuensis Ramose
Pseudobatostom
ella
igoi Ramose
Pseudobatostom
ella
kuramotoi Ramose
Pseudobatostom
ella
sp. Ramose
Capitanian Baten beni Zid Series (Lower
Biohermal Complex/Dolomies
Inferieures)
Cyclotrypa sp. Encrusting
Capitanian Capitan Fm Fenestella sp. Fenestrate
Goniocladia sp. Fenestrate
Polypora sp. Fenestrate
Capitanian Chandalaz Fm Epiactinotrypa sp. Encrusting
Etherella crassa Fenestrate
Hinganotrypa conspecta Fenestrate
Girtypora clara Ramose
Girtypora regula Ramose
Tavayzopora septata Ramose
Girtypora sp. Ramose
Hayasakapora sp. Ramose
262
Age Formation Genus Species Morphology
Tavayzopora sp. Ramose
Ulrichotrypella sp. Ramose
Capitanian Gerster Limestone Meekopora angularis Bifoliate
Araxopora paraxensis Dendroid
Dyscritella bogatensis Dendroid
Dyscritella incrustata Dendroid, Encrusting
Dyscritella indenta Dendroid, Encrusting
Dyscritella irregularis Dendroid, Encrusting
Dyscritella multistyla Dendroid, Encrusting
Dyscritella obtusa Dendroid
Dyscritella paratenuimuralis Dendroid
Dyscritella rotundis Dendroid, Encrusting
Dyscritella tenuimuralis Dendroid
Dyscritellina acuta Dendroid
Dyscritellina macrostyla Dendroid
Hinganella augusta Dendroid
Hinganella polygona Dendroid
Hinganella regularis Dendroid
Morozoviella sp. A Dendroid
Morozoviella sp. B Dendroid
Morozoviella sp. C Dendroid
Neoeridotrypella petita Dendroid
Neoeridotrypella plana Dendroid
Paralioclema cf. nekhoroshevi Dendroid
Paralioclema grandis Dendroid
Paralioclema multiphragma Dendroid
Paralioclema neospinigera Dendroid
Pseudobatostom
ella
frondosa Dendroid
263
Age Formation Genus Species Morphology
Rhombotrypella malforma Dendroid
Stellahexiforma diaphragma Dendroid
Stellahexiforma gerterensis Dendroid
Stenodiscus annulata Dendroid, Encrusting
Stenodiscus delicata Dendroid, Encrusting
Stenopora abnormis Dendroid
Stenopora curriensis Dendroid
Stenopora dyscritellensis Dendroid
Stenopora malvanensis Dendroid
Stenopora microautoformis Dendroid
Stenopora multiacanthensis Dendroid
Dyscritellina pseudoclivosa Encrusting
Fistulipora elegantula Encrusting
Fistulipora monticulosa Encrusting
Stenopora thuliformis Encrusting
Rectifenestella cf. microretiformis Fenestrate
Fenestella fragilis Fenestrate
Kingopora inflata Fenestrate
Kingopora wardlawi Fenestrate
Polypora kasanensis Fenestrate
Polypora keyserlingi Fenestrate
Polypora keyserlingiformis Fenestrate
Polypora soyanensis Fenestrate
Polyporella helgersoni Fenestrate
Rectifenestella cordiretiformis Fenestrate
Wjatkella cf. permiana Fenestrate
Wjatkella hemiseptifera Fenestrate
Wjatkella problemus Fenestrate
Wjatkella wjakensis Fenestrate
Timanotrypa sp. A Laminar
264
Age Formation Genus Species Morphology
Cyclotrypa multiformis Massive, Encrusting
Cyclotrypa ogbinensis Massive, Encrusting
Fistulipora dybowskii Massive, Encrusting
Ulrichotrypella multitubulosa Ramose
Timanodictya sp. B Ramose
Timanodictya sp. C Ramose
Timanodictya sp. D Ramose
Reteporidra depressa Reticulate
Capitanian Iwaizaki Fm - Unit 8 Coscinotrypa? sp. Bifoliate
Meekopora densa Bifoliate
Ramipora ambigua Bifoliate, Ramose
Goniocladia intricata Fenestrate
Fistulipora cf. timorensis Encrusting
Pseudobatostom
ella
yamazakii Erect
Penniretepora akiyamai Fenestrate
Penniretepora iwaizakiensis Fenestrate
Penniretepora rectodichotoma Fenestrate
Penniretepora tenuis Fenestrate
Coeloclemis minima Lamellate
Dyscritella cylindrica Ramose
Dyscritella iwaizakiensis Ramose
Hayasakapora erectoradiata Ramose
Pseudobatostom
ella
igoi Ramose
Pseudobatostom
ella
microstoma Ramose
Capitanian Maokou Fm Girtypora sp. Ramose
265
Age Formation Genus Species Morphology
Hayasakapora sp. Ramose
Timanodictya zunyiensis Ramose
Capitanian Merbah el Oussif Series -
Djebel Tebaga Biohermal
Complex
Cyclotrypa sp. Encrusting
Capitanian Oskhtinskaya Fm Permolioclema iraidae Encrusting
Hayasakapora gracilis Erect
Polypora hinganensis Fenestrate
Polypora inaudita Fenestrate
Clausotrypa costata Ramose
Dyscritella bobilevi Ramose
Dyscritella praespinigera Ramose
Dyscritella tchurkensis Ramose
Parastenodiscus hardmaniformis Ramose
Primorella polita Ramose
Streblascopora punctata Ramose
Capitanian Hexagonella tortuosa Bifoliate, Ramose
Araxopora araxensis Erect
Polypora tubulosa Fenestrate
Surmaq Fm - Unit 3
Septopora tarazi Fenestrate
Capitanian Whitehorse Sandstone Fm Leioclema dozierense Ramose
Capitanian Xiangbo reef Dybowskiella sp. Encrusting
Fistulipora sp. Encrusting
Fenestella donaica Fenestrate
Fenestella sp. Fenestrate
266
Age Formation Genus Species Morphology
Polypora guangxiensis Fenestrate
Polypora sp Fenestrate
Ascopora cf. quadritubulata Ramose
Eridopora sp Ramose, Encrusting
Rhabdomeson sp. Ramose
Capitanian Yates-Tansill Fm Girtypora sp. Ramose
Capitanian Zewan Fm Dyscritella cf. iwaizakiensis Erect
Dyscritella sparsigemmata Erect
Stenopora ? kashmirensis Erect
Polypora transiens Fenestrate
Acanthocladia anceps Pinnate
Hayasakapora sp. Ramose
Wuchiapingian Ford Fm Corynotrypa voigtiana Encrusting
Fenestella retiformis Fenestrate
Kingopora ehrenbergi Fenestrate
Synocladia virgulacea Fenestrate
Acanthocladia sp. Pinnate
Dyscritella columnaris Ramose, Encrusting
Wuchiapingian Fistulipora elegantula Erect
Hambast Fm - Unit 6
Fistulipora pseudomonticulosa Erect
Wuchiapingian Qubuerga Fm Dyscritella bogatenis Ramose
Dyscritella cf. lucida Cylindrical
Maychellina ornata Cylindrical
Rhombopora sp. B Cylindrical
Araxopora? cf. bifoliata Frondose
267
Age Formation Genus Species Morphology
Wuchiapingian Zechstein Fm Dyscritella tenuimuralis Encrusting, Dendroid
Ulrichotrypa incrustata Encrusting
Dyscritella angularis Encrusting, Dendroid
Dyscritella microstoma Encrusting
Dyscritella tubulosa Encrusting
Kavariella typica Erect
Penniretepora waltheri Erect
Acanthocladia f]diffusa Erect
Acanthocladia laxa Fenestrate
Kingopora ehrenbergi Fenestrate
Spinofenestella geinitzi Fenestrate
Spinofenestella minuta Fenestrate
Thamniscus perplexus Fenestrate
Fenestella sp. Fenestrate
Rectifenestella retiformis Fenestrate
Ryhopora delicata Fenestrate
Synocladia virgulacea Fenestrate
Kingopora baderi Fenestrate
Acanthocladia anceps Pinnate
Acanthocladia minor Pinnate
Penniretepora waltheri nodata Pinnate
Wuchiapingian Zewan Fm Dyscritella tenuirama Erect
Changhsingian Chhidru Fm Septopora sp. Fenestrate
Synocladia sp. Fenestrate
Acanthocladia sp. Pinnate
Changhsingian Upper Changxing Fm - Tudiya Fistulipora sp. Encrusting
268
Age Formation Genus Species Morphology
Acanthocladia cf. quadalupensis Ramose
Buildup
Rhombopora? sp. Ramose
Changhsingian Zewan Fm Dyscritella tenuirama Erect
Stenodiscus cf. chaetetiformis Erect
Hayasakapora grossa Ramose
Abstract (if available)
Abstract
Changes in the diversity and environmental distribution of marine stenolaemate bryozoans through the Permian to Early Jurassic interval are used to constrain the timing of environmental stress, determine the degree to which it differentially affected marine settings, evaluate its long-term impact on bryozoan evolution, and review current extinction scenarios for the end-Permian and end-Triassic crises. Results indicate that the end-Permian and end-Triassic mass extinctions were characterized by protracted intervals of environmental stress that initiated during the end-Guadalupian event several million years prior to the end-Permian, and again during the Norian stage prior to the end-Triassic. Environmental degradation lasted into the Early Triassic and Early Jurassic, hampering the recovery of marine communities. Prior to both extinctions offshore settings were affected first, suggesting environmental stress resulted from the gradual encroachment of a deep-water phenomenon onto the shelves, thus precluding catastrophic events as kill mechanisms and supporting long-term oceanographic processes such as widespread euxinia resulting from massive volcanism and global warming. The end-Permian event was the most influential for bryozoans, triggering a permanent change in their paleoenvironmental preferences from onshore to mid-shelf settings and a taxonomic switch between stenolaemate and gymnolaemate bryozoans. The taxonomic switch itself initiated a gradual shift in the overall morphological composition of bryozoan assemblages between erect-dominated Paleozoic communities and encruster-dominated Mesozoic communities. Major changes in the paleoenvironmental preferences of bryozoans were also coupled with fluctuations in their diversity.
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University of Southern California Dissertations and Theses
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Asset Metadata
Creator
Powers, Catherine Marie
(author)
Core Title
Unraveling mass extinctions: Permian to Early Jurassic onshore-offshore trends of marine stenolaemate bryozoans
School
College of Letters, Arts and Sciences
Degree
Doctor of Philosophy
Degree Program
Geological Sciences
Publication Date
09/30/2009
Defense Date
07/07/2009
Publisher
University of Southern California
(original),
University of Southern California. Libraries
(digital)
Tag
Brachiopoda,Bryozoa,diversity,mass extinctions,morphology,OAI-PMH Harvest,paleoecology,paleoenvironments,Permian-Triassic,Triassic-Jurassic
Language
English
Contributor
Electronically uploaded by the author
(provenance)
Advisor
Bottjer, David J. (
committee chair
), Corsetti, Frank A. (
committee member
), Edmands, Suzanne (
committee member
)
Creator Email
jamet@usc.edu,lespowers@mac.com
Permanent Link (DOI)
https://doi.org/10.25549/usctheses-m2629
Unique identifier
UC1493487
Identifier
etd-Powers-3188 (filename),usctheses-m40 (legacy collection record id),usctheses-c127-256152 (legacy record id),usctheses-m2629 (legacy record id)
Legacy Identifier
etd-Powers-3188.pdf
Dmrecord
256152
Document Type
Dissertation
Rights
Powers, Catherine Marie
Type
texts
Source
University of Southern California
(contributing entity),
University of Southern California Dissertations and Theses
(collection)
Repository Name
Libraries, University of Southern California
Repository Location
Los Angeles, California
Repository Email
cisadmin@lib.usc.edu
Tags
Brachiopoda
Bryozoa
mass extinctions
morphology
paleoecology
paleoenvironments
Permian-Triassic
Triassic-Jurassic