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Journal of Paleontology: LOWER MISSISSIPPIAN TRILOBITE B...http://findarticles.com/p/articles/mi_qa3790/is_200707/ai_n194...1 of 98/12/2008 12:35 AM
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Brezinski, David K  ABSTRACT-Six stratigraphically distinct trilobite faunas are recognized in the Lower Mississippian strata of the central United States. These faunas range inage from earliest Kinderhookian to Meramecian, and are, in ascending order: Pudoproetus missouriensis, Comptonaspis swallowi, Proetidesinsignis-Perexigupyge, Breviphillipsia semiteretis, Exochops portlockii, and Hesslerides bufo. Trilobite species diversity waxed and wanedthrough the early Mississippian of the central United States, but reached its maximum during the late Kinderhookian within the Comptonaspisswallowi fauna. The Comptonaspis swallowi, Breviphillipsia semiteretis, and Exochops portlockii faunas of the mid-continent can be correlated with the C. swallowi, B. semiteretis, and Hesslerides arcentensis faunas of the Caballero and Lake Valley Formations of New Mexico. The verticaldistribution, composition, and diversity variations among individual faunas suggest that they are evolutionarily discrete and therefore of  biostratigraphic utility. Their stratigraphic distribution appears to be controlled by sea level and climatic fluctuations.New Osagean trilobites identified and described are Exochops burlingtonensis n. sp. and Richterella carteri n. sp. from the Burlington Formationof Missouri, Australosutura osagensis n. sp. from the Keokuk Limestone of Missouri and the Arcente and Dona Ana Members of the Lake Valley Formation of New Mexico, and Spergenaspis boonensis n. sp. from the Boone Formation of Oklahoma.INTRODUCTIONTRILOBITES are relatively abundant and diverse from the Kinderhookian strata of the central United States. Conversely, Osagean species appearto be rarer, less diverse, and less well known, as indicated by the dearth of taxonomic papers discussing them. Two decades of collecting fromOsagean strata has confirmed to the author that the decrease in trilobite diversity from the Kinderhookian to the Osagean is not just an artifact of fewer published studies.That early Mississippian trilobites exhibit a marked diversification paralleling the transgressive phase of the Kaskaskia sequence has been noted by Brezinski (1986, 1999). This diversification is correlated with an early Mississippian adaptive radiation of many marine invertebrate groupsand is equivalent to Stage 1 of late Paleozoic trilobite evolution as proposed by Brezinski (1999). Although Stage 1 of Brezinski (1999, fig. 3) ischaracterized by a distinct group of genera, three specific faunas restricted to the Kinderhookian, early Osagean, and late Osagean, respectively, were also recognized. Further refinement of the stratigraphie ranges of trilobite species now allows recognition of six distinct faunas within early Mississippian strata of the central United States. These faunas illustrate a distinct biostratigraphic segregation of lower Mississippian trilobitespecies. This refined trilobite biostratigraphy for the Mississippian has evolutionary implications as well as stratigraphie utility.This paper describes six biostratigraphic faunas from the Lower Mississippian of the mid-continent region of the United States, and describes andillustrates new Osagean forms. All descriptive terminology is based on Whittington (1997), and new materials are reposited in the section of Invertebrate Paleontology at Carnegie Museum of Natural History (CM), Pittsburgh, Pennsylvania, the paleontology collections of the University of Missouri (UMC), and the University of Illinois (U of 111).BIOSTRATIGRAPHIC ZONATION OF LOWER MISSISSIPPIAN TRILOBITES Although taxonomic studies of Kinderhookian trilobites of the central United States are numerous (e.g., Hessler, 1962a, 1962b, 1963, 1965;Brezinski, 1986, 1988a, 1988b, 200Oa), it appears that much less attention has been paid to younger strata, as indicated by the paucity of published studies of that interval (see, for example, Hessler, 1965). A review of the literature on early Mississippian trilobites of the centralUnited States shows that, of the more than fifty species known, over forty are known only from the Kinderhookian, primarily the lateKinderhookian. The remaining ten species are from Osagean and Meramecian strata. While these numbers suggest that Kinderhookian speciesare more abundant, widespread, and diverse than their Osagean counterparts, it could be argued that the Osagean strata are simply less wellstudied. Indeed, studies of Osagean trilobites of the United States are exceedingly sparse (Hessler, 1965; Brezinski, 1986, 200Ob). However, aftersearching Osagean strata of the central United States for two decades, the author feels that stratigraphie variations in species diversity areprobably real rather than artifacts. Those field forays, in conjunction with an extensive search through museum and university collections,helped to more precisely define species ranges, and in turn verified the dramatic stratigraphie changes in trilobite species diversity in the centralUnited States. When these stratigraphie ranges are plotted, a distinct biostratigraphic zonation of trilobites is identifiable. Six separate faunas arerecognizable, and this zonation refines trilobite species stratigraphie distributions as previously delineated by Brezinski (1999).PUDOPROETUS MISSOURIENSIS FaunaThe earliest Mississippian trilobites known from the United States are holdovers from the end of the Devonian. Two species, Pudoproetusmissouriensis (Shumard, 1855), and Australosutura spinosus (Herrick, 1888), are known from the latest Devonian Louisiana Limestone of northeastern Missouri (Williams, 1944). Of these two species, only Pudoproetus missouriensis (=P. obesa Branson and Andrews, 1938) rangesthrough to the earliest Mississippian (early Kinderhookian) Glen Park Limestone and Bushburg Sandstone of eastern Missouri and the Prospect
Journal of Paleontology: LOWER MISSISSIPPIAN TRILOBITE B...http://findarticles.com/p/articles/mi_qa3790/is_200707/ai_n194...2 of 98/12/2008 12:35 AM
Hill Sandstone of Iowa (Fig. 1 ). This interval is equivalent to the Hannibal Shale, Horton Creek Formation, and Bachelor Sandstone of Missouri.This fauna spans the lower to middle Kinderhookian, Siphonodella sulcata to Siphonodella sandbergi conodont zones (Canis, 1968; Collinson et al.,1971).COMPTONASPIS SWALLOWI Fauna A stratigraphically younger and much better known trilobite fauna is present in upper Kinderhookian strata of the central United States. Speciesof this fauna are known from the Compton Limestone of Missouri and southwestern Illinois and the Rockford Limestone of Indiana. This fauna isthe most diverse (20 species) late Paleozoic trilobite fauna recognized in the United States (Brezinski, 1999, 200Ob). Termed fauna A of Stage 1 by Brezinski (1999), this faunal association is best known from the lower Chouteau Formation of central Missouri (Fig. 1). Herein this fauna isnamed the Comptonaspis swallowi fauna for the most common component. Other important faunal members include Ameropiltonia lauradanaeBrezinski, 2000, Elliptophillipsia ellipticus (Meek and Worthen, 1865), Griffithidella welleri (Branson and Andrews, 1938), Breviphillipsiasampsoni (Vogdes, 1887), and Dixiphopyge armatus (Vogdes, 1887) (Brezinski, 1988a). Ancillary trilobites of this fauna are Namuropyge sp,,Proetides colemani Messier, 1962, and Brachymetopus brezinskii Hahn and Hahn, 1996. This fauna is contemporary with the Siphonodellacrenulata and 5. isosticha conodont zones of the central United States (Canis, 1968; Collinson et al., 1971). Although characteristic of the centralUnited States, many genera of this fauna, including the eponymous species, are also present in the Caballero Formation of New Mexico (Brezinski,200Ob).PROETIDES INSIGNIS-PEREXIGUPYGE Fauna A late Kinderhookian and earliest Osagean association of trilobite species, included by Brezinski (1999) in fauna A, is present in the upperChouteau Formation of northeastern Missouri, the basal Sedalia Dolomite of central Missouri, and the Starr Cave and Gilmore City Formations of Iowa (Brezinski, 1988b, 200Oa). This fauna is characterized by Proetides insignis (Winchell, 1868), as well as a number of species of Perexigupyge, including P. chouteauensis Brezinski, 2000, P. hodgesi Brezinski, 1988, and P. gerki Brezinski, 1988 (Fig.l). Also present in thisfauna are Richterella snakedenensis Messier, 1965, Breviphillipsia trophis Messier, 1963, and Richterella hessleri Brezinski, 2000. Brezinski(1999) did not separate this fauna within his Stage 1 species, but later (Brezinski, 200Oa) recognized that it was distinct from the subjacent C.swallowi fauna. This fauna is present within the early Mississippian sequence of New Mexico, as indicated by the occurrence of an undeterminedspecies of Proetides Walter 1924 in the Andrecito Member of the Lake Valley Formation. This trilobite fauna is equivalent to the upperSiphonodella isosticha and perhaps also the Gnathodus punctatus conodont zones.BREVIPHILLIPSIA SEMITERETIS FaunaThe third distinct Mississippian trilobite fauna arose in the early Osagean, and is analogous to Brezinski's (1999) Stage 1, fauna B. This fauna ispresent in the Fern Glen Limestone of Missouri; the Burlington Limestone of Missouri, Illinois, and Iowa; and the Reed Spring Limestone of southwestern Missouri (Fig. 1). The components of this fauna include Breviphillipsia semiteretis Hessler, 1963, Piltonia tuberculata (Meek and Worthen, 1870), PiItonia brevicomus (Hessler, 1963), Griffithidella dons (Hall, 1860), Pudoproetus femglenensis (Weller, 1909), Exochops burlingtonensis n. sp., and Richterella carteri n. sp. It was also recognized by Brezinski (200Ob) within the Andrecito through Arcente Membersof the Lake Valley Formation of New Mexico. This trilobite fauna is equivalent to the Gnathodus punctatus through Scaliognathus anchoralisconodont zones.EXOCHOPS PORTLOCKII Fauna Within late Osagean strata, a widespread but low-diversity fauna can be recognized that is herein named the Exochops portlockii fauna. Thisfauna is equivalent to fauna C of Brezinski (1999) and is present within shelf deposits of the Keokuk and Warsaw Formations of Iowa, Missouri,and Illinois; the Edwardsville Formation of Indiana; the Fort Payne Formation of Kentucky; and the Boone Formation of Oklahoma (Fig. 1). TheH. arcentensis fauna from the Dona Ana Member of the Lake Valley Formation of New Mexico (Brezinski, 200Ob) is considered equivalent. Inaddition to E. portlockii (Meek and Worthen, 1865), this fauna contains Spergenaspis mauvaisensis (Hessler, 1965), Spergenaspis boonensis n.sp., Pudoproetus cf. P. femglenensis, Basidechenella timwhitei Leiberman, 1994, and Australosutura osagensis n. sp.The Exochops portlockii fauna is equivalent to the Gnathodus bulbosus and Gnathodus texanus conodont zones.HESSLERIDES BUFO FaunaOverlying the Exochops portlockii fauna is a depauperate fauna consisting of two species that are both geographically and stratigraphically restricted. These are Hesslerides bufo (Meek and Worthen, 1870) and Spergenaspis salemi Brezinski, 1987. This fauna is present within the upper Warsaw Formation of eastern Missouri; the Salem Limestone of Missouri, Illinois, and Indiana; and the St. Louis Limestone of Missouri (Fig. 1).Brezinski (1999) did not identify this as a separate fauna, but with more recent stratigraphie study (Brezinski, 200Ob), it can be recognized as adistinct association. Hesslerides pustulosus (Snider, 1915) from the Moorefield Formation of northeastern Oklahoma may represent an equivalentoff-shelf biofacies. However, this species is associated with Australosutura aff. A. gardneri (Ormiston, 1966) and an undescribed species of Phillibole Richter and Richter, 1937. Both of these genera show little biostratigraphic utility because of their long ranges even though they aregeographically widespread.DEPOSITIONAL SEQUENCES AND EARLY MISSISSIPPIAN TRILOBITE EVOLUTION IN THE CENTRAL UNITED STATES
Journal of Paleontology: LOWER MISSISSIPPIAN TRILOBITE B...http://findarticles.com/p/articles/mi_qa3790/is_200707/ai_n194...3 of 98/12/2008 12:35 AM
 While Brezinski (1999, fig. 3) was able to identify three specific lower Mississippian shelf faunas (A through C) in the central United States, thesix faunas identified herein greatly improves the previous resolution. The biostratigraphic delineation of the trilobite species described aboveappears to represent discrete shallow water associations. This discreteness and the wide disparity in species diversity between individual faunasappear to reflect environmental fluctuations (Brezinski, 1986; 200Oa).Comparison of the stratigraphie distribution of the trilobite faunas and the vertical distribution of depositional sequences suggests a high level of correspondence between faunal boundaries and sea level low stands, but this relationship is not one-to-one (Fig. 2). For example, theComptonaspis swallowi and Proetides insignis-Perexigupyge faunas are present within the same thirdorder sequence. However, theirstratigraphie and geographic distribution may be attributable to both environmental and temporal factors (Brezinski, 200Oa). Conversely, theBreviphillipsia semiteretis, Exochops meramecensis, and Hesslerides bufo faunas are vertically constrained by sequence boundaries. Thus,individual trilobite faunas appear to be partly, but not wholly, constrained by transgressive-regressive sequences.The low species diversity during the Pudoproetus missouriensis fauna and Hesslerides bufo fauna appears to bracket intervals when speciesdiversity waxed during the late Kinderhookian (Comptonaspis swallowi fauna) and then steadily dwindled through the Osagean into theMeramecian. Species diversity does not rebound until the late Mississippian (Brezinski, 1999). A contemporaneous diversity drop has been notedin the Arundian and Holkerian (= late Osagean to early Meramecian) of Great Britain (Thomas et al., 1984; Riley, 1993). It should be noted thatno similar trends in faunal stratigraphie segregation or diversity patterns are evident in deep-water deposits along the southern margin of theUnited States. Brezinski (1999) attributed this change in species diversity among shelf species to an early Mississippian transgress!ve episode(Kaskaskia transgression of Sloss, 1963). Consequently, the early Mississippian diversity changes appear to reflect global rather than regionalconditions. Although Pudoproetus Hessler, 1963, and Australosutura Campbell and Goldring, 1960 commonly occur in deep water deposits of Oklahoma,Texas, and Georgia (Rich, 1966; Brezinski, 1998), the only recognized occurrences of this association within shallow water deposits are in thelatest Devonian Louisiana Limestone (Pudoproetus missouriensis fauna) and the late Osagean (Exochops portlockii fauna). It appears that thesegenera had physiological tolerances that permitted existence in presumably cold and deep-water environments. The reemergence of these twogenera in shallow water faciès may signal a repetition of such environmental conditions during the latest Devonian and late Osaeean.Feist and Petersen (1995) interpreted Pudoproetus species as disaster taxa based on their global distribution following the Frasnian-Famennianextinction episodes (Hallam and Wignall, 1997). While the reasons for this Famennian extinction episode are open to debate, there is littlequestion that the latest Devonian is characterized by global cooling and high-latitude glaciation (Veevers and Powell, 1987; Copper, 1986, 1998;Isaacson et al., 1999). Therefore, the middle Mississippian reappearance of these two deep-water genera within shallow water deposits may reflectchanges in water conditions.The latest Devonian and late Osagean intervals are both characterized by global sea level lowering (Ross and Ross, 1988; Johnson et al., 1985), which appears to have been caused by brief but distinct glacial episodes (Veevers and Powell, 1987). Although these two periods of SouthernHemisphere glaciation are interpreted to have caused drops in sea level, their effect on global climate has only recently been identified (Isaacsonet al., 1999). Consequently, the reappearance of Pudoproetus and Australosutura (two characteristically deep water and presumably cold watergenera) within shallow water environments can be interpreted to reflect a repeated invasion of shelf environments by cold water during twoseparate periods of global cooling.In this light, the early Mississippian diversity pattern exhibited by trilobites of the central United States can be interpreted as being controlled by a period of global warming (and concurrent sea level rise) bracketed by episodes of global cooling and associated sea level drop. Therefore thelower Mississippian trilobite faunas of the central United States appear to be controlled by both climatic and third-order sea level sequences.SYSTEMATIC PALEONTOLOGY Family BRACHYMETOPIDAE Prantl and Pfibyl, 1950Genus AUSTRALOSUTURA Campbell and Goldring, in Amos et al., 1960 AUSTRALOSUTURA OSAGENSIS new speciesFigure 3.1-3.10 Australosutura sp. BREZINSKI, 2000b, p. 1049, fig. 5.18-5.20.Diagnosis.-Pygidium highly vaulted, strongly arched longitudinally. Axis broad, approximately 0.43 of maximum pygidial width, strongly arched, overhanging posterior margin. Pleurae not posteriorly recurved.Description.-Pygidium highly vaulted, narrow, with a subtriangular outline. Axis broad, robust, highly elevated, moderately posteriorly tapering, 0.43 times the total pygidial width, sharply rounded at terminus, semicircular in transverse profile with vertical sides especially tothe posterior, mildly arched in longitudinal profile with posterior terminus overhanging rear margin, composed of 12 robust rings with shallow, wide interring furrows. Rings ornamented with coarse tubercles. Pleural fields narrow, outline acutely triangular, strongly arched in transverseprofile, mildly arched in longitudinal profile, composed of seven straight ribs that extend beyond the margin into a short marginal spine. Eachrib comprised of a highly elevated anterior rib and narrow, lower posterior rib.

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