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Stasis and extinction of Silurian (Llandovery-Wenlock) trilobite associations related to oceanic cyclicity

Mikulic, Donald G

ABSTRACT-Silurian trilobites of the central United States belong to a series of temporally-successive associations which appeared abruptly,
maintained taxonomic stasis for a time, and then disappeared abruptly. Their disappearance resulted from global perturbations of short-term
duration and moderate magnitude, which caused substantial taxonomic replacement but no reorganization of major ecosystems. The most
significant extinction and replacement in Silurian trilobite associations in the study area occurs near the Llandovery-Wenlock boundary. This
turnover in trilobite associations appears to correspond to Jeppsson's Ireviken Event in his model of oceanic and climatic cyclicity. Major sea-level
changes earlier in the Llandovery did not have a similar impact on trilobite associations.

INTRODUCTION

A NUMBER of trilobite associations have been identified from the Silurian rocks of the central United States (Mikulic, 1979; in press), which show
distinct patterns of abrupt appearance, followed by taxonomic stasis, and then abrupt extinction (Kluessendorf and Mikulic, 1997). Some of these
associations are restricted to either a reef or nonreef setting; however, key taxa may range through a variety of environments, demonstrating
that the succession of associations is an evolutionary feature and not an environmental one. The best-documented of these associations are found
in the upper Llandovery-lower Wenlock rocks of this region where non-trilobite biostratigraphic control and depositional history are well
established. Within this time interval, evidence indicates that extinction events may be related to the Ireviken Event (Jeppsson, 1997), a
stressful transitional period between a primo and secundo oceanic episode (Jeppsson, 1990; Aldridge et al., 1993; Jeppsson et al., 1995; Jeppsson,
1998), as opposed to being related to specific sea level changes. This interval includes what is probably the most important global biotic
extinction event within the Silurian.

BACKGROUND

Beginning more than 140 years ago, a diverse assemblage of Silurian trilobites, which were among the first described from that time period in
North America, was collected and described from the central United States (e.g., Hall, 1867; Weller, 1907; Raymond, 1916). Unfortunately,
very little biostratigraphic, paleoecologic, or other data accompanied the resulting published descriptions or contemporary museum collections,
making it difficult to arrange these taxa into any kind of useful framework. Moreover, attempts at refining the systematics of these trilobites
have been of limited success because of the difficulty of collecting new material and the resulting dependence on poorlydocumented museum
specimens.

Over the last 30 years, however, we have made extensive new trilobite collections from this region and compiled a wide variety of associated
stratigraphic, biostratigraphic, geographic, paleoecologic, and other data. As a result, we have been able to place many of these taxa into a
precise biostratigraphic and environmental framework, which has enabled us to recognize certain patterns of extinction, stasis, and replacement
and to thereby gain important insights into the evolution and extinction of these trilobites.

SETTING

In the study area, which comprises the central United States (Illinois, Wisconsin, Michigan, Iowa, Indiana, Ohio), the Silurian rocks are
predominantly carbonates that were deposited in shallow epicontinental seas. While some stratigraphic units represent restricted environments,
most contain normal-marine biotas characterized by a diversity of benthic taxa. The area is especially well known for an abundance of
Wenlock-Ludlow reefs containing a high diversity of echinoderms, cephalopods, brachiopods, trilobites, and other organisms.

Silurian taxa described from this area were generally treated as a single biota in most early reports (e.g., Hall, 1867). In the early 20th century,
however, it was recognized that several distinct stratigraphic and geographic groupings of trilobite taxa might be present (Weller, 1907;
Raymond, 1916), but difficulty in establishing a reliable litho- and biostratigraphic framework for the region and the scarcity of
well-documented collections available for study greatly limited progress. Lowenstam's classic work in the 1940s and 1950s helped to establish
some of the environmental features of the biota, especially reef taxa (Lowenstam, 1948, 1957). Since the 1960s, new outcrop and subsurface
information have presented the opportunity to establish a reliable stratigraphic framework to restudy the trilobite biota.

Based on lithologic and biotic data, we are able to recognize two distinct paleogeographic regions in the study area that existed during the
Llandovery, possibly paralleling paleolatitude (Mikulic et al., 1996). The northern region includes northeastern Wisconsin, northwestern
Illinois, northern Michigan, and eastern Iowa, as well as much of Ontario and North Greenland. The southern region comprises southern
Michigan, southeastern Wisconsin, northeastern Illinois, Indiana, Ohio, and areas to the south and east. Depositional regimes during the
pre-Telychian coincide with this paleogeography: a thick section of peritidal and shallow subtidal carbonate strata, as well as
coral/stromatoporoid biostromal deposits, dominated the northern region (Ehlers and Kesling, 1957; Kluessendorf and Mikulic, 1989, 1994a;
Harris and Waldhuetter, 1996), whereas a thinner section of mostly carbonate sediments deposited under relatively deeper subtidal conditions
was present on the south (Mikulic et al., 1996; Watkins and Kuglitsch, 1997).

These pre-Telychian strata are truncated by a major unconformity and sequence boundary, which coincides with the contact between the

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historically-recognized Alexandrian and Niagaran series of the Silurian in this region (Mikulic, 1977, 1979; Kluessendorf and Mikulic, 1992,
1996a). This unconformity marks an episode of widespread emergence and karstification that followed a major eustatic sea level fall brought
about by glaciation in the southern hemisphere (Kluessendorf, 1990; Kluessendorf and Mikulic, 1996b, 1998). This break in deposition is
recognized in many other places in the world (Aldridge et al., 1993). On the basis of conodonts and brachiopods, none of the sub-unconformity
strata in the study area have been dated as younger than Aeronian (Kluessendorf and Mikulic, 1994a, 1996a; Mikulic and Kluessendorf, 1998).

The irregular topography at this unconformity was transgressed in the early to mid Telychian by argillaceous sediments, which are otherwise
uncharacteristic for the Silurian in the study area. In fact, this dramatic change from carbonate sediments below the unconformity to
argillaceous sediments above is recognized worldwide (Jeppsson, 1990). These sediments contain abundant and diverse conodonts, which indicate
that the transgression began as early as the Pterospathodus celloni Conodont Biozone and extended into the succeeding P. amorphognathoides
Conodont Biozone (Kleffner et al., 1994; Norby et al., 1996). Globally, the greatest diversity of Silurian conodonts occurs at this time (Jeppsson,
1990; Aldridge et al., 1993). The disappearance of the P. amorphognathoides Conodont Biozone occurs at about the Llandovery-Wenlock
boundary (TelychianSheinwoodian). Throughout this interval, sediments gradually became less argillaceous. Wenlock strata are dominantly
pure carbonates including bioclastic shoal, lagoonal, and subtidal shelf sediments. Reefs and carbonate banks began to develop and proliferate
during the Wenlock in the study area (Kluessendorf et al., 1995). Although conodonts are quite rare in these carbonate strata, the basal strata
fall within the Kockelella ranuliformis Conodont Biozone (Kleffner et al., 1995).

TRILOBITE ASSOCIATIONS

Trilobites are low in both abundance and diversity, and trilobite associations are poorly defined throughout much of the Llandovery in the study
area. This is probably related to the prevalence of marginal-marine conditions and possibly cooler temperatures caused by waning glaciation. It
is clear, however, that the illaenid Stenopareia and related genera are among the most common and environmentally widespread trilobites,
typifying the majority of strata in this interval.

During the Telychian, the conspicuous Stenopareia imperator-Ekwanoscutellum laphami trilobite association (Fig. 1), characterized by illaenids
(Stenopareia) and large scutelluines (Ekwanoscutellum, Meroperix, Opoa), became environmentally and geographically widespread.
Subordinate taxa in this association include encrinurines (Distyrax, Perryus), cheirurids (Chiozoan) and phacopids (Acernaspis). This association
was prevalent in a variety of both reef and level-bottom environments, including reefs in the Attawapiskat Formation of the Hudson Bay
Lowlands (Norford, 1981; Gass and Mikulic, 1982; Rudkin and Westrop, 1997), North Greenland (Lane, 1972, 1988; Lane and Peel, 1980), the
Brandon Bridge Formation of southeastern Wisconsin (Kluessendorf et al.,1995) and the Johns Creek Quarry Member of the Scotch Grove
Formation of eastern Iowa (Mikulic, 1979); and level-bottom environments in the Engadine Dolomite of Michigan and northeastern Wisconsin
(Kluessendorf and Mikulic, 1994a), the Brandon Bridge of southeastern Wisconsin and northeastern Illinois (Kluessendorf and Mikulic, 1994b),
and the Farmers Creek Member of the Hopkinton Formation in eastern Iowa (Mikulic, 1979). In much of this area, this association occurs in
strata that were deposited during the transgression across the major pre-Telychian unconformity. Stenopareia persisted through these major
Telychian sea level fluctuations.

The most significant extinction affecting the character of the Silurian trilobite fauna in the central U.S., when the dominant trilobite taxa
became extinct, occurs near the Llandovery-Wenlock (Telychian-Sheinwoodian) boundary (Fig. 2). This contrasts sharply with the limited
change in reef trilobite faunas at the Ordovician-Silurian boundary when no important trilobite taxa disappeared despite the severity of the
extinction event on the biota overall (Mikulic, 1981, 1985). The Stenopareia imperator-Ekwanoscutellum laphami trilobite association
disappeared at the end of the Pterospathodus amorphognathoides Conodont Biozone, and was replaced by the Bumastus ioxus-Kosovopeltis acamus
trilobite association (Mikulic, in press). The most conspicuous aspect of this turnover is the replacement of Stenopareia and related illaenid taxa
by a diversity of morphologically-similar bumastines, such as Bumastus. Only a single, rare illaenid, "Stenopareia" pterocephalus, continued
beyond this extinction event to co-exist for a short time with bumastines in the early Wenlock (Kluessendorf and Mikulic, 1994a). This turnover
occurs in both reef and nonreef environments, and the bumastine and scutelluine taxa occur in similar proportions to the illaenids and
scutelluines within each environment. Phacopids, cheirurids, and encrinurines, which remain subordinate taxa in the Wenlock, also show
similar patterns of change.

The Bumastus ioxus-Kosovopeltis acamus trilobite association is typical of reef and carbonate bank environments (Mikulic, 1979, in press;
Mikulic and Hickerson, 1994) (Figs. 3, 4), although it also occurs in some nonreef settings (Fig. 5). This association correlates with the Kockelella
ranuliformis Conodont Biozone at a time when prolific reef development was beginning in Wisconsin, Illinois, and Iowa. Although some of the
trilobite taxa are controlled environmentally and may be endemic to either a reef or nonreef setting, Bumastus ioxus and Kosovopeltis acamus
occur in both reef and nonreef environments.

Several other sequential trilobite associations can be recognized in the study area, however, poor nontrilobite biostratigraphic control precludes
their precise dating. For example, the reef and carbonate bank environments characterized by the Bumastus ioxus-Kosovopeltis acamus
association appear to be succeeded by lagoonal-subtidal shelf deposits containing the Sthenarocalymene celebra trilobite association (Fig. 6)
(Mikulic, in press) throughout most of the study area. This association is succeeded by the Glyptambon verrucosus-Calymene breviceps trilobite
association (Fig. 7) (Mikulic, in press), which is found in similar level-bottom environments. The Glyptambon verrucosus-Calymene breviceps
association is found in the Racine Dolomite in Illinois and in the Waldron Shale of Indiana and Tennessee. Interestingly, this association seems to
co-occur with different brachiopod communities in carbonate and argillaceous sediments. This replacement is also reflected in the reef
environment where key association taxa such as Glyptambon gassi, Litotix armata, and Cybantyx insignis occur (Fig. 8).

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STASIS

Silurian trilobite associations in the study area exhibit three levels of stasis. The first level of stasis in all of these trilobite associations is
short-term taxonomic stasis at the species level. Each association appears abruptly, maintains its taxonomic composition for some time, and then
disappears abruptly. This level is similar to the coordinated stasis described by Brett and Baird (1995) and Brett et al. (1996).

At a higher level of taxonomic stasis, trilobites in associations are mostly replaced by related genera. For example, within Wenlock reef
environments, bumastines always dominate in abundance and diversity, although they are represented by different genera and species in
successive communities (Mikulic 1979, 1981, in press). This level of taxonomic stasis is similar in scale to community stasis within Boucot's
(1983) ecologic evolutionary units.

The third type of stasis represents the long-term persistence of specific groups of morphologies in specific environments, which, in some cases, is
independent of phyletic origin. We term this morphotypic stasis. This level of stasis is demonstrated by trilobite taxa in reef and related
environments which exhibit a limited range of morphologies throughout most of the Paleozoic, from the Middle Ordovician to the end of the
Devonian (Mikulic, 1979, 1981). No significant additions are made to this group of morphologies over this extensive time interval, although
there are some important deletions. While individual taxa appear and disappear on various time scales, the overall composition of this
morphotypic association remains constant in terms of morphologies represented and their abundance and diversity. In a few cases,
phyletically-unrelated trilobites may represent the same morphologic position successively, with the replacement taxon mimicking its
predecessor's general features. Undoubtedly, this long-term morphotypic stasis represents a fundamental environmental stability that far
exceeds the range of most species and genera, and is more commonly observed on the level of families and orders. Morphotypic stasis operates on a
scale greater than Boucot's ecologic evolutionary units, and changes in these types of associations reflect fundamental environmental
reorganization on a global scale.

OCEANIC CYCLICITY

A model of oceanic and climatic cyclicity for the Silurian was proposed by Jeppsson (1990) and refined by Aldridge et al. (1993), Jeppsson et al.
(1995), and Jeppsson (1998). This model postulates that variations in carbon dioxide content between the atmosphere and oceans produced
alternating episodes of specific climatic and environmental conditions that can be related to certain sedimentological and biotic changes. It
further hypothesizes that biotic extinctions occurred during stressful periods of turnover, or events, in oceanic state between these episodes.
Jeppsson (1997) noted that the extinctions commonly took place in a stepwise manner, reflecting fluctuating conditions that repeatedly stressed
the faunas during this transition period. Moreover, Jeppsson (1990) proposed that habitat displacement accompanying sea-level fluctuations,
could not, by itself, explain the extinctions he observed. Among his evidence for this, Jeppsson noted that Chatterton et al. (1990) related the
appearances and disappearances of Llandovery and Wenlock trilobite and conodont clades in northwestern Canada to transgressive-regressive
events, yet their plots showed that, in some cases, the taxonomic turnovers preceded the interpreted change in sea level.

Jeppsson's model is controversial, and there is debate as to the timing of his episodes and events and their relation to sea level fluctuations (e.g.,
see Loydell, 1998; Johnson, 1996). However, his lithologic and biotic predictions agree very well with our observations of late Llandovery-early
Wenlock strata in the study area.

DISCUSSION

The replacement of the Stenopareia imperator-Ekwanoscutellum laphami trilobite association by the Bumastus ioxus-Kosovopeltis acamus
trilobite association does not correlate with the pre-Telychian regression, which marks the most significant sea level fall in the Silurian of the
region (and perhaps globally), even though this would normally be considered a severe environmental disruption capable of causing a major
extinction. Nor does it coincide with the onset of the subsequent transgression. Brett et al. (1996) also observed that turnover between intervals
of coordinated stasis did not necessarily correspond to sea level fluctuations or sequence boundaries in the Devonian of the Appalachian basin. In
our study area, this significant extinction and replacement of trilobite associations appears to correlate with the disappearance of the P.
amorphognathoides Conodont Biozone during the Ireviken Event (Aldridge et al., 1993; Jeppsson, 1997), a period of turnover between oceanic
states (Fig. 2). The Ireviken Event had the most far-reaching effects of all the events in Jeppsson's model, affecting many taxonomic groups
globally. For example, 80 percent of conodont taxa became extinct during this event, representing the most profound effect on the conodont
fauna during the entire Silurian (Jeppsson, 1997). Ramsk*ld (1985) observed that -50 percent of the trilobite fauna on Gotland became extinct
at this time also. Brett and Eckert (1989) also reported a significant late Llandovery pelmatozoan extinction falling within the Ireviken Event.
Acritarchs, ostracodes, graptolites, and foraminifers show a similar significant decline (Aldridge et al., 1993; Jeppsson, 1997). The fact that
"Stenopareia" pterocephalus persisted for a time to co-exist with Bumastus may reflect the stepwise extinction postulated to take place during
these events.

The replacement of the Sthenarocalymene celebra trilobite association by the Calymene breviceps-Glyptambon verrucosus trilobite association
may coincide with the late Wenlock (Homerian) Mulde Event (Jeppsson et al., 1995) (Fig. 2). The Mulde Event is marked by the most profound
extinction of Silurian graptolites (Jeppsson et al., 1995). An increase in argillaceous sedimentation occurs at this time globally, including the
study area.

Because trilobites are not particularly abundant and are absent from parts of the column in the study area, it is unlikely that the ranges for all
trilobite taxa can be plotted with the precision possible for conodonts. Therefore, all of the subtle changes identified in the conodont biota cannot

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be recognized in trilobite associations. It may still be possible, however, to establish the temporal relationships between many of these trilobite
associations and the episodes and events of Jeppsson's model based on non-conodont criteria such as lithologic patterns. This will be particularly
useful in conodont-poor strata which are common throughout the study area.

If trilobite associations can be linked to Jeppsson's model of oceanic cyclicity, it is then possible that taxonomic turnover in associations results
from short-term global environmental perturbations which lack the capacity to reorganize ecosystems. In contrast, the disappearance of the
long-term morphotypic associations result from profound global environmental perturbations that yield a permanent reorganization of an
ecosystem.

CONCLUSIONS

Understanding the paleogeographic, paleoecologic, and biostratigraphic framework of specific taxa can help to explain evolutionary changes in
trilobite associations. Using such a framework, we have been able to recognize that the most significant turnover in Silurian trilobite associations
of the central United States is at the Llandovery-Wenlock boundary.

At this time, the dominant trilobite taxa in both reef and nonreef settings became extinct and were replaced by related taxa, mostly at the
generic level, and from within the same morphotypic association. Detailed biostratigraphic data have allowed us to establish a correlation
between this trilobite faunal turnover and the Ireviken Event, a global environmental perturbation, identified by Jeppsson et al. (1995) and
Jeppsson (1997). A preceding major sea level change, for which there is ample physical evidence in the study area, had little impact on the
composition of trilobite associations.

Correlation of trilobite extinction and environmental events suggests that trilobite associations in the study area can be used as biostratigraphic
tools, which could be especially useful in sediments where the benthic biota is diverse and abundant but biostratigraphically-useful microfossils
are rare.

Additional, although less pronounced, extinctions of Silurian trilobite associations occurred in the study area, but their timing is less precise
because verifying biostratigraphic data are presently unavailable. Preliminary evidence suggests that these additional extinctions may coincide
with other events and episodes in Jeppsson's model.

ACKNOWLEDGMENTS

We acknowledge the numerous quarry operators who gave us access to their properties and museum personnel who gave us access to their
collections. We extend special thanks to R. Norby and M. Kleffner for processing and identifying conodonts used to construct the biostratigraphic
framework. Much of the biostratigraphic work was funded by National Science Foundation grant DEB 92-01048 to the authors. We also thank M.
Johnson and an anonymous reviewer for their comments on the manuscript. Published with permission of the Chief of the Illinois State Geological
Survey.

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ACCEPTED 23 OCTOBER

DONALD G. MIKULIC AND JOANNE KLUESSENDORF

Illinois State Geological Survey, Champaign, Illinois 61820, , and Department of Geology, University of Illinois, Urbana, Illinois 61801,

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