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rise and fall of late Paleozic trilobites of the United States, The

Brezinski, David K

ABSTRACT-Based on range data and generic composition, four stages of evolution are recognized for late Paleozoic trilobites of the contiguous
United States. Stage 1 occurs in the Lower Mississippian (Kinderhookian-Osagean) and is characterized by a generically diverse association of
short-ranging, stenotopic species that are strongly provincial. Stage 2 species are present in the Upper Mississippian and consist of a single,
eurytopic, pandemic genus, Paladin. Species of Stage 2 are much longer-ranging than those of Stage 1, and some species may have persisted for as
long as 12 m.y. Stage 3 is present within Pennsylvwian and Lower Permian strata and consists initially of the eurytopic, endemic genera Sevillia
and Ameura as well as the pandemic genus Ditomopyge. During the middle Pennsylvanian the very long-ranging species Ameura missouriensis
and Ditomopyge scitula survived for more than 20 m.y. During the late Pennsylvanian and early Permian, a number of pandemic genera
appear to have immigrated into what is now North America. Stage 4 is restricted to the Upper Permian (late Leonardian-Guadalupian) strata
and is characterized by short-ranging, stenotopic, provincial genera.

The main causal factor controlling the four-stage evolution of late Paleozoic trilobites of the United States is interpreted to be eustacy. Whereas
Stage 1 represents an adaptive radiation developed during the Lower Mississippian inundation of North America by the Kaskaskia Sequence,
Stage 2 is present in strata deposited during the regression of the Kaskaskia sea. Stage 3 was formed during the transgression and stillstand of the
Absaroka Sequence and, although initially endemic, Stage 3 faunas are strongly pandemic in the end when oceanic circulation patterns were at
a maximum. A mid-Leonardian sea-level drop caused the extinction of Stage 3 fauna. Sealevel rise near the end of the Leonardian and into the
Guadalupian created an adaptive radiation of stentopic species of Stage 4 that quickly became extinct with the latest Permian regression.

INTRODUCTION

ALTHOUGH THE Carboniferous and Permian represent nearly a third of the total history of the class Trilobita (Owens and Hahn, 1993), little
attention has been paid to the evolutionary patterns within this time interval. Those studies that have discussed the evolutionary trends for
trilobites within the late Paleozoic have dealt mainly with intrageneric morphological trends (Weller, 1937; Hahn and Hahn, 1967;
Chamberlain, 1969). Even though such studies are important, morphological trends are the result, not the cause, of evolutionary episodes.

Some of the more recent studies have illustrated the progressive decline of generic diversity through the late Paleozoic (Eldredge, 1978, fig. 9;
Foote,1993, Table 2), which has led some authors to consider Permo-Carboniferous trilobites as incidental components of the benthic
communities (Boucot, 1983), or attributed their diminishing generic numbers to the rise of durophagus predation (Signor and Brett, 1984).
Although these hypotheses may be partially valid, for trilobites they are based upon outdated data. Figure 1 illustrates variations in the number
of trilobite genera through the Permo-Carboniferous. Figure 1.1 presents data from Foote (1993, table 2) and Figure 1.2 is a pictorial
representation of generic (including subgenera) diversity compiled for this study from more than 40 publications that date back to about 1970.
Although the sources are too voluminous to cite here individually, they represent the more recent contributions on late Paleozoic trilobite
taxonomy from Europe, Asia, Australia, and North America.

The two illustrations in Figure 1 indicate that within the Lower Carboniferous there is more than a 300-percent discrepancy between the two
data sets. While this discrepancy decreases to almost 200 percent in the Upper Carboniferous, it reaches nearly 300 percent again in the Upper
Permian. Comparison between these data with trilobite generic diversity for the middle Paleozoic (Foote, 1993, table 2) indicates that the Lower
Carbonif erous has a greater number of trilobite genera than any comparable interval of time after the Ordovician. This interpretation is based
on the presumption (likely false) that the number of recognized Devonian and Silurian trilobite genera has not increased since the origination of
Foote's data set. However, any attempt to make inferences about trilobite evolution based on antiquated data (such as the 1959 Treatise on
trilobites) is tenuous at best.

The global variations in trilobite generic diversity trends illustrated in Figure 1.2 parallels the generic diversity pattern observed for the
Carboniferous and Permian of the United States (Fig. 2). The pattern begins with a peak in generic and species diversity in the Lower
Mississippian, continues with decreasing numbers in both categories in the middle Carboniferous, then proceeds with relatively steady but small
numbers in the Upper Carboniferous and Lower Permian, and finally closes with an increase in both genera and species in the Upper Permian.

The goal of this paper is to discuss generic diversity and species range patterns of Carboniferous and Permian trilobites of the United States and to
discuss their paleoecologic and paleogeographic distributions. These new data provide insight into the tempo and mode of late Paleozoic trilobite
evolution.

Figured type specimens are reposited at the U.S. National Museum (USNM), Field Museum of Natural History, Chicago (FMNH UC), Carnegie
Museum of Natural History (CM), University of Missouri (UM), University of Illinois (UI), University of Wisconsin (UW), Museum of Northern
Arizona (MNA), and the Illinois Geological Survey (IGS).

STRATIGRAPHIC DISTRIBUTION

Although Chamberlain (1969, text-fig. 4) attempted to illustrate the ranges of North American Carboniferous trilobite species, his diagram

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tended to lump species and ranges so that a picture of their overall stratigraphic ranges is unclear. Figure 2 is a composite range chart of species
of Carboniferous and Permian trilobites from the contiguous United States. Data illustrated in this figure are taken from a wide range of
publications as well as from ongoing studies by the author. The time scale (third column from left) and the relative position and duration of the
geologic stages (second column from left) are extrapolated from Cowie and Bassett (1989).

Stratigraphic range durations of trilobite species tend to vary systematically through time. Ranges during the early Mississippian, on average,
are short, varying from 2 to 5 m.y. Late Mississippian species durations are considerably longer and can, as in the case of Paladin chesterensis, be
as long as 12 m.y. During the Pennsylvanian and early Permian (Wolfcampian), species durations are very long. For instance, Ameura
missouriensis ranged through most of the Pennsylvanian and into the early Permian, approximately 30 m.y. Other species such as Ditomopyge
scitula and Ditomopyge decurtata have estimated ranges of 20 and 15 m.y, respectively. During the late Permian, species ranges again tended to
become short, averaging 2 to 5 m.y.

The above-mentioned variations in species durations as well as generic compositional changes permit subdivision of the range chart into four
stages. Although further subdivision based on species composition is possible, the four-fold subdivision was utilized, since each stage, as currently
recognized, is generically distinct. Stage 1 encompasses the early Mississippian (Kinderhookian-early Meramecian); Stage 2 is found in the late
Mississippian (Meramecian-Chesterian) and earliest Pennsylvanian; Stage 3 in the Pennsylvanian and early Permian (late Morrowanearly
Leonardian); and finally Stage 4 appears in the late Permian (late Leonardian-Guadalupian). To understand the causal factors that produced this
distribution it is necessary to address individual species in each stage.

STAGE 1

The initial stage of late Paleozoic trilobite evolution in the United States occurs within Lower Mississippian strata. These faunas are well known,
having been described by Hessler (1963, 1965), Chamberlain (1969), and Brezinski (1986b, 1988a, 1988b, 1988c, 1998). This is the most
generically diverse stage, with 17 genera currently known from the Kinderhookian through early Meramecian strata of the United States.
Figure 3 illustrates the ranges of some of the more widespread species found in this stage.

Composition.-Characteristic genera of Stage 1 include: Pudoproetus, Breviphillipsia, Griffithidella, Comptonaspis, Piltonia, Richterella,
Dixiphopyge, Spergenaspis, Exochops, and Griffithides (Fig. 4). Species of this stage are generally short-ranging and are typically restricted to
one of three recognized faunas (Fig. 3). Fauna A is found in middle to late Kinderhookian strata. They include the Cuyahoga Formation of the
Appalachians, the Chouteau Formation of Missouri, and the Caballero Formation of New Mexico. Characteristic species of fauna A include
Breviphillipsia sampsoni, Griffithidella welleri, Comptonaspis swallowi, Dixiphopyge armata, and species of Brachymetopus as well as a number
of less widely distributed species. Fauna B is present in the Burlington Formation of Missouri, lower Lake Valley Formation of New Mexico, and the
Woodhurst Member of the Lodgepole Formation of Montana. Species that characterize this fauna include Breviphillipsia semiteretis, Griffithidella
doris, Piltonia tuberculata, and Proetides insignis. The youngest fauna in Stage 1, fauna C, is present in the Warsaw through Salem Formations of
the Midcontinent, and the upper Lake Valley Formation of New Mexico. Characteristic components are Exochops portlocki (Meek and Worthen),
several species of Gri,ffithides, and the genus Spergenaspis.

Paleoecology.-Brezinski (1986b) discussed the distribution of trilobite species from the Chouteau Formation (Kinderhookian) of central Missouri.
He proposed that several species, such as Comptonaspis swallowi, Dixiphopyge armata, Elliptophillipsia ellipticus, and Proetides colemani, were
found in a restricted set of lithologies, suggesting that these species were limited to a narrow range of environmental settings. This led to the
interpretation that such species were stenotopic. Similarly, Brezinski (1987, 1988b) found that species of Spergenaspis and Perexigupyge were
found almost solely in oolitic and bioclastic lime grainstones, suggesting the optimal habitat of these species included high-energy shoal-water
conditions. While most genera were confined to shallow shelf settings, some genera (e.g., Australosutura, Carbonocoryphe, Phillibole, and
Thigriffides) were restricted to deeper environments of the outer shelf, shelf-edge, and starved oceanic basin (Brezinski, 1990, 1998). This
segregation of genera from shallow water to deep water environments suggests a biofacies arrangement for Lower Carboniferous trilobites of the
United States similar to that seen by Hahn and Hahn (1988) in the Tournaisian Kolenkalks of Belgium.

The paleoecological distribution of genera in Stage 1 seems to indicate that constituents of this stage tended to be stenotopic. While some genera
can be found in a broad range of environmental settings and are probably eurytopic, such as Griffithidella, the majority of genera are confined to
a narrow range of lithologies and thus environmental settings. Moreover, species of this stage appear to be consistently associated with relatively
stenotopic epifaunas, such as corals, brachiopods, and crinoids, rather than typical biofacies of infaunal molluscs which tend to dominate late
Paleozoic nearshore environments (Anderson, 1971).

Paleogeography.-The Lower Mississippian shallow- to deep-water biofacies mentioned above and discussed by Brezinski (1990; 1998) indicates a
generic segregation of trilobites reminiscent of that in the Lower Paleozoic (see for example Fortey, 1975, and McNamera and Fordham, 1981).
Lower Mississippian trilobites, like those of the Lower Paleozoic, include highly provincial shelf genera and cosmopolitan off-shelf genera.

Of the 17 genera known from Stage 1, only Brachymetopus, Namuropyge, and Piltonia have been reported from shelf strata outside of the United
States. Eleven genera found exclusively in shelf strata are native to North America (e.g., Breviphillipsia, Comptonaspis, Richterella, Exochops,
Dixiphopyge, and Spergenaspis). Those genera found in the shelf-edge and deep-ramp environments, such as Pudoproetus, Phillibole, and
Australosutura, are known worldwide from strata of this time interval. Likewise, the deep-water, starved ocean basin fauna of Carbonocoryphye
and Phillibole are known to have a wide geographic distribution.

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Figure 5 illustrates the stratigraphic variation in diversity and endemism of late Paleozoic trilobites of the United States. The greatest number of
genera of the late Paleozoic of the United States occurs in the Lower Mississippian. Of the 17 genera currently recognized, nearly 75 percent are
endemic to the United States. Sando and others (1975) found a similar provincialism among Lower Carboniferous corals of the central United
States.

STAGE 2

In contrast to the high generic diversity observed in Stage 1, Stage 2 displays the lowest trilobite generic diversity of the four stages. Whereas
Stage 1 boasts 17 genera, Stage 2 contains a single genus. This genus, Paladin, is a geographically widespread genus with much longer-ranging
species than Stage 1. Although Paladin first occurs in the Osagean (Cisne, 1967), it is an extremely rare component of Stage 1. While most other
genera that comprise Stage 1 become extinct by the early to middle Meramecian, Paladin survived this early Mississippian extinction episode to
become the dominant trilobite constituent in the late Meramecian and Chesterian.

Composition.-The genus Paladin has more species than any other late Paleozoic trilobite genus known from North America. Paladin has been an
enigmatic generic designation since its erection by Weller (1936). In its original designation Paladin was listed as a late Mississippian genus that
lacked a frontal border; whereas Kaskia was a similar genus that possessed an anterior border. Whittington (1954) found that the presence and
absence of an anterior border varied within species during ontogeny and placed Kaskia as a subgenus of Paladin. Cisne (1967) proposed that the
two genera were synonyms, a point with which Osmolska (1970) agreed. Hahn and Hahn (1970), on the other hand, also considered Kaskia to be
a subgenus of Paladin. Much of the uncertainty surrounding the generic designation of Kaskia and Paladin appears to be the result of the broad
range of morphologies exhibited by species of Paladin (Fig. 6).

As many as 15 species of Paladin are currently recognized within late Mississippian strata of the United States. Unfortunately, many of these
species are based on single specimens and as a result may simply represent aberrant phenotypes of a very few species. Because of the morphologic
breadth exhibited by species of this genus, it is plausible that some of these species may not be valid.

The species of Stage 2 tend to be much longer-ranging than species of Stage 1 (Fig. 7). For example, P. chesterensis ranges from the late
Meramecian throughout most of the Chesterian, a duration of as much as 12 m.y. Other species such as P. girtyianus and P. rosei range through
most of the Chesterian, a duration of perhaps 7 to 10 m.y. Paladin morrowensis is present in late Chesterian and earliest Morrowan strata, a
species duration of probably 5 to 7 m.y. In summary, Stage 2 contains a fauna with low generic diversity and long-ranging, phenotypically
plastic species.

Paleoecology.-Species of Paladin tend to occur in many different lithologies, indicating a broad range of depositional environments. Detailed
examination of the distribution of Paladin chesterensis occurrences in late Mississippian strata of the Appalachians indicates a strong association
of this trilobite species with certain biofacies (Brezinski, 1983a). Brezinski found that P. chesterensis occurred more than 70 percent of the time
with nuculid bivalve biofacies that were present in nearshore depositional environments. This species also was found within more offshore, mixed
bivalve-brachiopod biofacies (20 percent of the occurrences), and to a much lesser degree with open shelf, brachiopod biofacies.

A similar pattern occurs in P. girtyianus. This species is most often found associated with molluscan faunas of the Fayetteville Shale of Arkansas
(Gordon, 1969), and to a much lesser degree with the brachiopod-crinoid biofacies of the Pitkin Limestone. At least two other species of Paladin, P.
moorei and P. longispina, are associated with nearshore molluscan biofacies. At this time it is not possible to draw any conclusions about the
paleoecologic distribution of any of the other species of this genus in North America, inasmuch as they have yet to be studied.

Consequently, based on these observations it is proposed that species of Paladin in Stage 2 tended to prefer nearshore habitats and are commonly
associated with molluscan biofacies. This preference for what is a typically unstable, if not harsh, environmental setting indicates that many of
the trilobite species of Stage 2 may have been eurytopic.

Paleogeography.-Paladin is one of the most stratigraphically and geographically widespread genera of the late Paleozoic (Osmolska, 1970). In
Europe, species of this genus are known from Great Britain, Poland, Spitzbergen, Germany, and Belgium (Osmolska, 1970; Hahn and Hahn,
1970). Paladin has also been recovered from Russia and Japan (Kobayashi and Hamada, 1980), as well as being recognized throughout North
America.

Owens and Hahn (1993) recognized two trilobite realms for this time interval. The United States is characterized by the Paladin-Sevillia Realm
and Eurasia by the Paladin-Cummingella Realm. Inasmuch as Sevillia is included in this paper within the overlying stage (Stage 3), and as most
of the genera that characterize the Eurasian Realm (i.e., Brachymetopus, Phillibole, and Pseudowarible; see Owens and Hahn, 1993, table 3) are
cosmopolitan through much of their stratigraphic range, it appears that for trilobites of the late Mississippian there is a total absence of
provincialism. Consequently, on a worldwide scale, this time interval is distinguished by a small number of pandemic genera.

Stage 2, which ranges from the late Meramecian to earliest Morrowan, is dominated by a single eurytopic, pandemic genus. In examining Figure
5, one should note how a large number of endemic genera of Stage 1 become extinct by the middle Mississippian (Meramecian), followed by the
proliferation of species of a single pandemic genus Paladin.

STAGE 3

Composition.-The long-ranging species of the genus Paladin of Stage 2 appear to give way to a number of new genera whose species are very

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long-ranging in Stage 3. These genera include: Sevillia, Ameura, and Ditomopyge (Fig. 8). These three genera arose in the early Pennsylvanian
(Morrowan-Atokan). Both species of Sevillia, S. trinucleata and S. sevillensis, appear to arise in the Morrowan to early Atokan and range to early
Desmoinesian, a duration of about 10 m.y. These two species are termed fauna A in Figure 9. During the Atokan, two additional species
originated; they ranged for a protracted period of time through most of the Pennsylvanian. One of these species, Ameura missouriensis, has a
species duration of as much as 30 m.y., from late Atokan until its last appearance in the Wolfcampian. The absolute range of the second species,
Ditomopyge scitula, is approximately 20 (Brezinski, 1988c; Brezinski et al., 1989) to 30 m.y. (Pabian and Fagerstrom, 1972, text-fig. 1).
Ameura missouriensis and D. scitula, whose combined ranges cover most of the middle Pennsylvanian, comprise what is here termed fauna B of
Figure 9. In the early Virgilian, a second species of Ditomopyge appears. This species, D. decurtata, ranges through the Virgilian and most of the
Wolfcampian, a species duration of between 12 and 15 m.y. Within this same interval, several genera, known from single occurrences, make
appearances in the United States. These include Brachymetopus (Acutimetopus) kansasensis, a species of Cheiropyge, Pseudophillispia sp., and
three species of Triproetus (Brezinski, 1992). All of these genera are better known from outside of North America during this time interval, and
constitute the exotic fauna C.

The genera that characterize the long-ranging Stage 3 fauna become extinct in the late Wolfcampian to early Leonardian. It is apparent that
these long-ranging species display a considerable period of stasis varying from 15 to 30 m.y.

Paleoecology.-The paleoecologic distribution of Pennsylvanian trilobites has been briefly discussed in a number of places (Bucurel and
Chamberlain, 1977; Brezinski, 1984, 1988a; Brezinski et al., 1989). The trilobite occurrences tend to recur with certain benthic biofacies
patterns in an onshore to offshore environmental gradient (see Stevens, 1971; Brezinski, 1983b). Table 1 illustrates occurrences of common
trilobite species of Stage 3 and their relationship to benthic biofacies. Each occurrence in this table represents a horizon or locality that has
yielded trilobite specimens. This table seems to indicate that common trilobite species of Stage 3 tended to occur within unstable, presumably
nearshore environments quite often within molluscan biofacies. This suggests that these species were eurytopic. Additionally, D. scitula has been
interpreted as an opportunistic species (Brezinski, 1986c) that occurred in large numbers along a single widespread stratigraphic horizon.
Opportunism is consistent with eurytopic habits.

Brezinski et al. (1989, fig. 3) have illustrated a trend of progressive size decrease for D. scitula in Atokan though late Missourian strata of eastern
Ohio. Recent findings from other locations in the United States further elucidates this trend. Figure 10 compares means of cranidial and pygidial
sizes for stratigraphically distinct populations of D. scitula. One can see that the means of populations from older stratigraphic intervals are
larger than the means from younger intervals. Weller (1935) illustrated a similar trend for length of the genal spines of Ditomopyge in
Pennsylvanian strata of the midcontinent region. Inasmuch as D. scitula is overwhelmingly found in nearshore biofacies (Table 1) and as no
apparent variation or trend in environmental preference is currently recognized, the reduction in size of this species is interpreted to be phyletic
rather than ecophenotypic. Such a temporally-based morphocline is suggestive of paedomorphism (McNamera, 1982), whereby ecological
pressures select for progressively smaller individuals. Unquestionably, this size reduction trend is worthy of further examination. The
interpretation that D. scitula was an opportunistic, eurytopic species that inhabited nearshore environments may indicate that the temporal
trend toward smaller-sized individuals represents a directional selection for smaller individuals in the unpredictable nearshore environments.

Although not fully documented, D. decurtata appears to exhibit an opposite size change from D. scitula. Specimens of D. decurtata from the
Virgilian are generally smaller and exhibit fewer ribs and rings than do individuals that occur in the Wolfcampian (Brezinski, 1988c, p. 942).
Recalling that D. decurtata tends to occur with more offshore biofacies than does D. scitula (see Table 1), it can be interpreted that the former
species tended to be more stenotopic, and inhabited more stable, offshore environments. Thus, the general trend toward size increase in D.
decurtata seems to indicate a selection toward increased size within a stenotopic species in stable environmental settings.

Paleogeography.-Initially, this stage is characterized by the appearance of three new genera (Sevillia, Ameura, and Ditomopyge). Two of these
genera, Sevillia and Ameura, are endemic (fauna A). Such generic endemism characterized the Morrowan (Ross and Ross, 1985a). Ditomopyge, a
pandemic genus, is known worldwide from Upper Carboniferous and Lower Permian strata. With the disappearance of both species of Sevillia in
the early Desmoinesian, only Ameura missouriensis and Ditomopyge scitula (fauna B) persist through much of the remainder of the
Pennsylvanian (Westphalian). During the Virgilian (late Stephanian) and Wolfcampian, Ditomopyge decurtata and a number of exotic
pandemic genera appear in the United States (i.e., Brachymetopyus (Acutimetopus), Pseudophillipsia, Cheiropyge, and Triproetus). The changes
from a largely endemic fauna (fauna A) during the early part of Stage 3 to a strongly pandemic fauna (fauna C) near the end of Stage 3 is
graphically represented in Figure 5.

Owens and Hahn (1993) indicate that throughout the Westphalian and Stephanian (middle and upper Pennsylvanian) only two trilobite realms
can be recognized worldwide. These realms are differentiated by numerous endemic genera, several of which (Brachymetopus, Paladin, and
Pseudophillipsia), are only endemic for this time interval, and could easily be considered cosmopolitan genera. Many of the truly endemic genera
(Sevillia, Ameura, Weania, Blodgettia, Alaskalethe) arose during the earliest part of Stage 3. Consequently, during much of this time interval
(Desmoinesian through Wolfcampian) provincialism is weak and many of the genera are cosmopolitan.

STAGE 4

Composition.-The long-ranging genera that characterize Stage 3 become extinct during the latest Wolfcampian to early Leonardian. During the
late Leonardian, a group of new genera appear and range through the end of the Guadalupian. Characteristic genera include Anisopyge, Delaria,
Vidria, and Novoameura (Fig. 11). The majority of Stage 4 species appear in the late Leonardian and early Guadalupian (Fig. 12). Species of this

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stage tend to be much shorter-ranging than those of the preceding stage with durations of less than 2 m.y. to as long as 7 m.y.

Paleoecology.-The majority of late Permian species are known from shallow-water facies of West Texas (Brezinski, 1992). These species tend to be
found within the platform-edge environments that surrounded the Midland, Delaware, and Marfa Basins. The species associated with these
environments are interpreted to be stenotopic, since they are associated with stable, brachiopod-echinoderm-bryozoan faunas (Newell and others,
1972, fig. 83).

A number of different species are present within the shelf areas that extended northwest from the West Texas basins. These species are
geographically widespread, and are known from the San Andres Formation of New Mexico, the Kaibab Limestone of Arizona, and the Phosphoria
Formation of Wyoming (Chamberlain, 1970; Cisne, 1971; Brezinski, 1991). Cisne (1971) concluded that most of the species from the late
Permian Kaibab Limestone of Arizona were eurytopes.

In summary, late Permian trilobites of the West Texas Permian Basin appear to be stenotopic species, inhabiting shelf-edge reefal facies. Those
species that inhabited shallow-water shelf environments of New Mexico, Arizona, and Wyoming were more eurytopic species, and tended to be
associated with molluscan biofacies.

Paleogeography.-The fauna of Stage 4 represents the most strongly endemic stage for genera of the late Paleozoic of the United States. Owens
(1983, text-fig. 1) has shown that the number of new genera that arose during the late Permian worldwide was notable, and that many of these
genera were endemic.

Owens and Hahn (1993) have proposed that two trilobite realms are discernable in the late Permian. The trilobite realms proposed by Owens and
Hahn (1993) for the Permian differ slightly from what is presented here because those authors divided the Lower and Upper Permian at a
different point than was done in this report. The division utilized here places the top of the Lower Permian at the top of the Roadian, well within
the Kungurian, and not at the top of that unit as shown by Owens and Hahn (1993, text-fig. 1). As a result of this difference in placement of the
stage boundary, genera such as Anisopyge and Delaria (and presumably Novoameura) are included in the realms of Owens and Hahn for the
Lower Permian as well as the Upper Permian, whereas they are considered restricted to the Upper Permian in this report.

DISCUSSION AND INTERPRETATION

The four stages of trilobite evolution within the late Paleozoic of North America reflect numerous controlling factors. Owens and Hahn (1993)
have proposed that the provincialism exhibited by trilobites of this interval was largely controlled by latitude. Although this likely was
important in their distribution, it is herein proposed that the main factor controlling the stratigraphic, geographic, and paleoecologic
distribution of late Paleozoic trilobites of the United States was the magnitude and frequency of sea-level fluctuations.

Figure 13 compares the 4 stages of trilobite development within the Permo-Carboniferous with second-, third-, and perhaps fourth-order sea-level
events of the late Paleozoic. Secondorder events are taken from Vail et al. (1977); the third- and/or fourth-order events are redrawn from Ross
and Ross (1988). The general trend for sea-level events for the Carboniferous and Permian reflect two major (second-order) events for this
interval. The earlier occurred in the Lower Carboniferous and can be equated with the Kaskaskia Sequence of Sloss (1963). The later event
spanning the Upper Carboniferous and Permian is equivalent to the Absaroka Sequence. Less extensive deepening events, which are
superimposed on the sequence-level deepenings, are evident within the Lower and Upper Mississippian, Pennsylvanian-Lower Permian, and
Upper Permian (see sea-level curve of Vail et al., 1977). The stages of trilobite ranges can ostensibly be correlated with these individual events.

Stage 1 of late Paleozoic trilobite evolution in North America occurred during the early Mississippian (Kinderhookian-Meramecian). This time
interval is characterized by a worldwide deepening episode that presumably began in the latest Devonian. This early Carboniferous transgression
follows a Late Devonian regression (Hangenberg Event) during which the last of the phacopid genera, that characterize the Devonian, became
extinct (Feist, 1991; Feist and Petersen, 1995). Following the Hangenberg Event, only a very few lazarus taxa, such as Pudoproetus, survived
(Feist and Petersen, 1995).

While the North American Midcontinent was submerged during the Kinderhookian, a wide range of habitats became available. Trilobites appear
to have quickly occupied and diversified into these habitats. This adaptive radiation produced a rapid increase in generic diversity, not only in
North America, but worldwide, during the Kinderhookian (Tournaisian) (Figs. 1, 5). The quick adaptation of trilobite genera to a wide range of
habitats is indicated by their tendency toward stenotopy during this interval of time (Brezinski, 1986a). The short ranges exhibited by many
species of this stage are reflective of the stenotopic tendencies of this fauna.

While overall sea level continued to deepen throughout the Kinderhookian and into the Osagean, a number of short-lived regressions punctuated
the stratigraphic record (Ross and Ross, 1985b, 1988). These events produced the species-level differentiation of the three individual faunas
recognized within this stage (Fig. 3). These three faunas appear to be contained within deposits of three separate sea-level events of either the
third- or fourth-order, and are characterized by species-specific faunas. The Kaskaskia Sequence reached its maximum depth during the Osagean.
However, generic diversity dropped dramatically immediately following this maximum, and nearly all shelf genera found in fauna A and fauna
B became extinct, leaving a new but impoverished fauna C. The rapid demise of the genera of faunas A and B is attributed to the "perched fauna"
phenomenon as discussed by Johnson (1974). Johnson proposed that a rapid sea-level drop near a transgressive maximum, where stenotopic
faunas are adapted to stable environmental conditions, leads to a concomitant extinction. Initiation of the Kaskaskia regression during the latest
Osagean and Meramecian brought Stage 1 to an end with the extinction of its characteristic genera of earlier faunas and appearance of the
pandemic genus Griffthides within the shelf faunas.

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Owens and Hahn (1993) have shown that three well-defined trilobite provinces characterize the Tournaisian (Kinderhookian, early Osagean) of
the world. Each provincial realm contains a strongly endemic faunal component. In the United States endemism is largely restricted to the shelf
faunas. Those genera found in off-shelf environments are commonly cosmopolitan (e.g., Phillibole, Carbonocoryphe, Australosutura, and
Pudoproetus). This provincialism is perplexing, given that the Lower Carboniferous is characterized by cosmopolitanism (Boucot, 1983, fig. 3),
and a panequatorial seaway with unrestricted circulation that existed between North America and Europe during this interval of time (Ross and
Ross, 1985a). Because of this apparent lack of a land barrier, Owens and Hahn (1993) suggested that the provincialism exhibited by the trilobite
faunas of this time was controlled by latitudinal variations. However, the strong provincialism is exhibited mainly by shelf genera. Genera that
characterize deep water environments (Phillibole, Carbonocoryphe) are cosmopolitan. Even though the distribution of genera may have also
been controlled by latitude, this was probably a minor contributor to the currently recognized distributional pattern.

The results of this study indicate that the provincialism displayed by trilobites of this interval is the result of a rapid adaptive radiation, where
the shelf genera arose and diversified locally, presumably from deep-water survivors of the Late Devonian extinctions. The absence of a land
barrier was inconsequential inasmuch as the stenotopic, shallow-water endemics could not migrate across the intervening oceanic barrier.

Species of the eurytopic genus Paladin, which characterize Stage 2, display long ranges and wide geographic and ecologic distributions. The
differences in species durations between Stages 1 and 2 are especially significant when one considers that the late Mississippian is characterized
by rapid sea-level oscillations (Sloss, 1963; Swann, 1964; Ramsbottom, 1979; Brezinski, 1989), and an overall reduction in ecospace due to the
withdrawal of the Kaskaskia sea.

Species of Paladin in the United States tended to be ecologic generalists (eurytopic). This may have aided in the trend for relative stasis of this
lineage, as well as its geographic pervasiveness (Lieberman, 1993). Jackson (1974) has shown that eurytopic species of bivalves tend to be more
cosmopolitan and have less tendency to speciate than stenotopic species. He attributed this to greater larval dispersal within eurytopic species,
since such forms tended to occur within more discontinuous shallow-water environments. The proliferation of species of Paladin in the late
Mississippian appears to have resulted from this eurytopic genus spreading worldwide during the regressive phase of the Kaskaskia Sequence.
Even though ecospace may have been diminishing during this time interval, individual species continued to survive unhindered.

The correlation between sea-level nadir at the end of the Mississippian and the end of Stage 2 is not precise. One species (P. morrowensis) can be
found in the earliest Pennsylvanian strata and represents the last representative of Stage 2. As sea level fluctuated during this low-stand (see
curve according to Ross and Ross, 1988), this presumed eurytopic species was able to exist in the greatly reduced ecospace. The components of
Stage 3 do not appear until somewhat later in the Morrowan. Sevillia appears in the Morrowan followed shortly thereafter by Ameura and
Ditomopyge. While Sevillia does not persist beyond the early Desmoinesian, the other two genera display extremely long ranges. Indeed, the
exceptionally long ranges of both A. missouriensis and D. scitula, in excess of 20 million years, is herein attributed to the eurytopic nature of both
of these species. The tendency of Ditomopyge scitula to associate with nearshore molluscan faunas, is interpreted to have produced a phyletic size
decrease and an affinity toward opportunism.

The appearence of a modest number of genera worldwide during the early Pennsylvanian suggests that there was a minor radiation of trilobite
genera concomitant with the deepening of the Pennsylvanian (Absaroka) sea. Any provincialism exhibited by early Stage 3 genera disintegrates
during the middle to late Pennsylvanian and early Permian when a number of pandemic genera appear in the central and southwestern United
States. Consequently, by the end of Stage 3 the faunas are characterized by pandemism. The pandemic fauna at this time is perplexing since the
United States, with the formation of Pangea and the uplift of the Appalachian-Ouachita-Marathon mountain belt, was geographically isolated
from Eurasia (Ross and Ross, 1985a). However, this period of geographic isolation also coincides with maximum deepening of the Absaroka sea
(Fig. 13). Therefore, dispersal of trilobite larvae may have been governed less by geographic barriers than by existing oceanic currents created
by maximum submergence during the Absaroka Sequence.

A marked regression in the early to middle Leonardian is correlated with the end of Stage 3 and the extinction of all existing trilobite genera
known in North America (Vail et al.,1977; Brezinski, 1992). The only areas that appear to have remained submerged during this low-stand of
sea level were deeper shelf and basinal settings of West Texas.

Within the middle to late Leonardian several new genera appear and represent the initiation of Stage 4, a very strongly provincial fauna. The
first appearance of these new genera is correlated with a deepening episode that began in the late Leonardian and continued into the Guadalupian
(Vail et al., 1977). The origination of the genera of Stage 4 during this deepening episode is equated to the adaptive radiation of trilobites seen in
the Kinderhookian (Stage 1). However, the late Permian transgression submerged much less shelf area than did the Lower Mississippian
deepening.

Owens (1983) has shown that the Upper Permian represents a period of radiation of trilobite genera worldwide. While Owens attributes this
diversification to the spread of reefs during this time, this study proposes that the increased diversifications and endemism are the result of this
late Permian deepening and concomitant adaptive radiation that produced a stenotopic fauna.

While there are many hypotheses as to the cause of the endPermian extinction, the emphasis of this paper is the extinction of trilobites from the
United States. Hypotheses implicating salinity (Fischer, 1964) or temperature (Stanley, 1984; Crowley and North, 1988) fall outside of the scope
of the current paper. However, Schopf (1974) and Simberloff (1974) have shown that reducing the area of shallow seas could have had a similar
effect on the number of families. Schopf (1974) attributed sea-level changes that affected the size of shelf areas near the end of the Permian to
variations in sea-floor spreading. Bilal et al. (1987, fig. 5) and Ross and Ross (1988, fig. 9) identified a marked drop in sea-level that corresponds

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to the end of the Guadalupian (Kazanian). This regression, which coincides with the end of the Capitanian, resulted in the destruction of
essentially all shelf ecospace in North America. Insofar as Stage 4 species are interpreted to be largely stenotopic, the destruction of habitable
environments would have ostensibly led to the extinction of shelf faunas. Therefore, the drop in sea level following the Guadalupian is interpreted
to have caused a "perched fauna" extinction (Johnson, 1974). The only other option, migration into deeper water environments, was precluded
since the Midland and Delaware basins had become silled and sites of evaporite deposition. Consequently, the extinction of trilobite genera within
the late Permian was unlike preceding evolutionary episodes in that there were no deep-water refuges to support survivors of the shelf
extinctions.

SUMMARY AND CONCLUSIONS

Figure 14 summarizes the interpretation presented herein. During the early Mississippian, transgression and the submergence of North America
made available abundant ecospace that led to an adaptive radiation of trilobites. This fauna was stenotopic and strongly endemic. Brief
shallowing episodes produced at least three separate faunas. Regional regression produced extinction of this perched fauna during the late
Osagean and Meramecian. Stage 2 developed in the late Mississippian during an overall period of regression. The trilobite fauna, composed of the
pandemic genus Paladin, tended to have species that were long-ranging and eurytopic. Stage 3 occurred during the Pennsylvanian and Lower
Permian. Although initially composed of endemic genera, the fauna became increasingly cosmopolitan by the end of Stage 3. Ranges of many
species of this stage were very long. The genera tended to be eurytopic and cosmopolitan. Stage 3 ended in the early to middle Leonardian with a
sharp regressive episode. Transgression during the late Leonardian and Guadalupian produced the strongly endemic, stenotopic fauna of Stage 4
in a brief, marked adaptive radiation. With regression at the end of the Permian, shelf ecospace was destroyed. No longer could trilobites emigrate
into basinal settings as these basins ultimately became sites of evaporite deposition. This led to the extinction of trilobites in the United States.

ACKNOWLEDGMENTS

This manuscript was greatly improved by helpful suggestions offered by C. A. Kertis and H. B. Rollins. Constructive comments were also provided
by reviewers B. S. Lieberman and R. Ludvigsen. The Maryland Geological Survey provided photographic facilities.

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DAVID K. BREZINSKI

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