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LOWER MISSISSIPPIAN TRILOBITE BIOSTRATIGRAPHY OF THE CENTRAL UNITED STATES, AND SOME NEW OSAGEAN
SPECIES

Brezinski, David K

ABSTRACT-

Six stratigraphically distinct trilobite faunas are recognized in the Lower Mississippian strata of the central United States. These faunas range in
age from earliest Kinderhookian to Meramecian, and are, in ascending order: Pudoproetus missouriensis, Comptonaspis swallowi, Proetides
insignis-Perexigupyge, Breviphillipsia semiteretis, Exochops portlockii, and Hesslerides bufo. Trilobite species diversity waxed and waned
through the early Mississippian of the central United States, but reached its maximum during the late Kinderhookian within the Comptonaspis
swallowi 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 vertical
distribution, 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 Formation
of 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.

INTRODUCTION

TRILOBITES are relatively abundant and diverse from the Kinderhookian strata of the central United States. Conversely, Osagean species appear
to be rarer, less diverse, and less well known, as indicated by the dearth of taxonomic papers discussing them. Two decades of collecting from
Osagean 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 groups
and is equivalent to Stage 1 of late Paleozoic trilobite evolution as proposed by Brezinski (1999). Although Stage 1 of Brezinski (1999, fig. 3) is
characterized 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 trilobite
species. 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 and
illustrates 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 central
United States shows that, of the more than fifty species known, over forty are known only from the Kinderhookian, primarily the late
Kinderhookian. The remaining ten species are from Osagean and Meramecian strata. While these numbers suggest that Kinderhookian species
are more abundant, widespread, and diverse than their Osagean counterparts, it could be argued that the Osagean strata are simply less well
studied. Indeed, studies of Osagean trilobites of the United States are exceedingly sparse (Hessler, 1965; Brezinski, 1986, 200Ob). However, after
searching Osagean strata of the central United States for two decades, the author feels that stratigraphie variations in species diversity are
probably 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 central
United States. When these stratigraphie ranges are plotted, a distinct biostratigraphic zonation of trilobites is identifiable. Six separate faunas are
recognizable, and this zonation refines trilobite species stratigraphie distributions as previously delineated by Brezinski (1999).

PUDOPROETUS MISSOURIENSIS Fauna

The earliest Mississippian trilobites known from the United States are holdovers from the end of the Devonian. Two species, Pudoproetus
missouriensis (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) ranges
through to the earliest Mississippian (early Kinderhookian) Glen Park Limestone and Bushburg Sandstone of eastern Missouri and the Prospect

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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. Species
of this fauna are known from the Compton Limestone of Missouri and southwestern Illinois and the Rockford Limestone of Indiana. This fauna is
the 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 is
named the Comptonaspis swallowi fauna for the most common component. Other important faunal members include Ameropiltonia lauradanae
Brezinski, 2000, Elliptophillipsia ellipticus (Meek and Worthen, 1865), Griffithidella welleri (Branson and Andrews, 1938), Breviphillipsia
sampsoni (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 Siphonodella
crenulata and 5. isosticha conodont zones of the central United States (Canis, 1968; Collinson et al., 1971). Although characteristic of the central
United 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 upper
Chouteau 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 this
fauna 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 undetermined
species of Proetides Walter 1924 in the Andrecito Member of the Lake Valley Formation. This trilobite fauna is equivalent to the upper
Siphonodella isosticha and perhaps also the Gnathodus punctatus conodont zones.

BREVIPHILLIPSIA SEMITERETIS Fauna

The third distinct Mississippian trilobite fauna arose in the early Osagean, and is analogous to Brezinski's (1999) Stage 1, fauna B. This fauna is
present 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 Members
of the Lake Valley Formation of New Mexico. This trilobite fauna is equivalent to the Gnathodus punctatus through Scaliognathus anchoralis
conodont 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. This
fauna 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). The
H. arcentensis fauna from the Dona Ana Member of the Lake Valley Formation of New Mexico (Brezinski, 200Ob) is considered equivalent. In
addition 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 Fauna

Overlying 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 a
distinct association. Hesslerides pustulosus (Snider, 1915) from the Moorefield Formation of northeastern Oklahoma may represent an equivalent
off-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 are
geographically widespread.

DEPOSITIONAL SEQUENCES AND EARLY MISSISSIPPIAN TRILOBITE EVOLUTION IN THE CENTRAL UNITED STATES

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While Brezinski (1999, fig. 3) was able to identify three specific lower Mississippian shelf faunas (A through C) in the central United States, the
six faunas identified herein greatly improves the previous resolution. The biostratigraphic delineation of the trilobite species described above
appears to represent discrete shallow water associations. This discreteness and the wide disparity in species diversity between individual faunas
appear 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, the
Comptonaspis swallowi and Proetides insignis-Perexigupyge faunas are present within the same thirdorder sequence. However, their
stratigraphie and geographic distribution may be attributable to both environmental and temporal factors (Brezinski, 200Oa). Conversely, the
Breviphillipsia 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 species
diversity waxed during the late Kinderhookian (Comptonaspis swallowi fauna) and then steadily dwindled through the Osagean into the
Meramecian. Species diversity does not rebound until the late Mississippian (Brezinski, 1999). A contemporaneous diversity drop has been noted
in the Arundian and Holkerian (= late Osagean to early Meramecian) of Great Britain (Thomas et al., 1984; Riley, 1993). It should be noted that
no similar trends in faunal stratigraphie segregation or diversity patterns are evident in deep-water deposits along the southern margin of the
United 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 regional
conditions.

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 the
latest Devonian Louisiana Limestone (Pudoproetus missouriensis fauna) and the late Osagean (Exochops portlockii fauna). It appears that these
genera had physiological tolerances that permitted existence in presumably cold and deep-water environments. The reemergence of these two
genera 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-Famennian
extinction episodes (Hallam and Wignall, 1997). While the reasons for this Famennian extinction episode are open to debate, there is little
question 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 reflect
changes 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 Southern
Hemisphere glaciation are interpreted to have caused drops in sea level, their effect on global climate has only recently been identified (Isaacson
et al., 1999). Consequently, the reappearance of Pudoproetus and Australosutura (two characteristically deep water and presumably cold water
genera) within shallow water environments can be interpreted to reflect a repeated invasion of shelf environments by cold water during two
separate 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 the
lower 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, 1950

Genus AUSTRALOSUTURA Campbell and Goldring, in Amos et al., 1960

AUSTRALOSUTURA OSAGENSIS new species

Figure 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 to
the 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 transverse
profile, mildly arched in longitudinal profile, composed of seven straight ribs that extend beyond the margin into a short marginal spine. Each
rib comprised of a highly elevated anterior rib and narrow, lower posterior rib.

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Etymology.-Named for the Osagean Series of the Mississippian, from which it is known.

Types.-Holotype, a pygidium from the Keokuk Limestone (Peerless Park Member) at Peerless Park, St. Louis County, Missouri, CM 53901.
Paratypes, 2 partial pygidia from the Keokuk Limestone (Peerless Park Member), same location as the holotype, CM 53902, 53903.

Additional material.-Two partial pygidia from the Arcente Member of the Lake Valley Formation, Otero County, New Mexico, CM 45737.

Distribution.-Present in the Keokuk Limestone of eastern Missouri and upper Arcente Member of the Lake Valley Formation, Otero County, New
Mexico.

Discussion.-The specimens of Australosutura recovered from the Peerless Park Member of the Keokuk Limestone of Missouri are identical to
specimens assigned to Australosutura sp. by Brezinski (200Ob) from the Arcente Member of the Lake Valley Formation of New Mexico.
Furthermore, the Australosutura specimens collected from Lake Valley Formation and the Keokuk species appear to be nearly coeval.
Australosutura osagensis n. sp. differs from other North American species of Australosutura by the stronger vaulting. It differs from A. llanoensis
Brezinski, 1999 by having a broader, more highly vaulted axis and narrower, more strongly arched transverse profile to the pleural fields. Since
these characters do not match well with any named North American species of this genus, the new species A. osagensis is herein erected.

The vaulting exhibited by Australosutura osagensis n. sp. is most similar to that of A. gardneri (Mitchell 1922) the type species of this genus, but
A. gardneri exhibits strongly posteriorly recurved pleurae and a narrower pygidial axis (0.37-0.39, compared to 0.43 for A. osagensis).

Family PHILUPSIIDAE Oehlert, 1886

Genus EXOCHOPS Weller, 1936

Discussion.-Exochops is a late Osagean genus that is characterized by elevated palpebrae, a bulbous glabella that reaches the anterior cranidial
margin, and an elongate, semielliptical pygidium. It is very similar to, and often mistaken for, the slightly younger genus Hesslerides except
that the latter genus has dorsally flattened palpebrae, slightly enlarged Ll, and a shorter, semicircular pygidium with a broader border. Both
genera display a restricted geographic distribution in the central and southern United States.

In erecting Exochops, Weller (1936) noted that this genus was similar to, and appeared to have been derived from, Griffithides Portlock, 1843.
The middle Visean and lower Namurian Eurasian genus Griffithides is younger than the late Tournaisian to early Viséan genus Exochops,
making Welter's proposed pattern of direct decent less likely. Hahn and Hahn (1969) and Hahn and Brauckmann (1991) proposed that
Griffithidinae was derived from the middle to late Tournaisian genus Griffithidella doris, a relatively common and widespread species from the
central and southern United States. Moreover, there are numerous morphological characters shared by Griffithidella doris and both Exochops
and Hesslerides (Table 1). These morphologic characters are the inflated glabella; reduced Ll; posterior located, enlarged palpebrae; broad
subocular groove; narrow straight occipital furrow; strongly shouldered, broad pygidial axis; and a bordered pygidium onto which the
interpleural furrows extend. The common morphologies and shared geographic distribution of Griffithidella, Exochops, and Hesslerides suggest
that these genera are part of a monophyletic group. This likelihood has prompted a phylogenetic evaluation of the known representatives of these
genera. For this evaluation Griffithidella doris was chosen as the outgroup in phylogenetic analysis and is the presumed ancestor. This
interpretation is consistent with Hahn and Hahn (1967) who used G. doris as the presumed ancestor in constructing a stratophenetic tree for
Griffithides. The list of 20 characters interpreted to be of phylogenetic significance were analyzed used character states given in Table 2. An
exhaustive search was conducted in PAUP 4.0 (beta version) using accelerated transformation optimization. From this analysis a single tree of 24
steps in length was retained. The single most parsimonious tree has a consistency index (CI) of 0.83 and is given in Figure 4. The topology
demonstrates that the two sister groups Exochops and Hesslerides are valid higher taxa that are temporally segregated. With the exception of the
displayed relationship between E. portlockii and E. burlingtonensis, there is complete congruence between the stratigraphic distribution of the
species and their grouping on the cladogram.

The species of Hesslerides are consistently aligned with their stratigraphie distribution. This interpreted monophyly demonstrates the inferred
close relationship and endemic nature of Exochops and Hesslerides. The demonstrable morphological similarity and interpreted monophyly
exhibited by these North American genera suggest further phylogenetic study of the Griffithidinae is necessary in order to ascertain the
relationship of these genera with what have been interpreted to be closely related European genera (e.g., Griffithides, Particeps Reed 1943, and
Cyphinoides Reed 1942) (Hahn and Brauckmann, 1991).

EXOCHOPS PORTLOCKH (Meek and Worthen, 1865) Figure 3.11-3.13

Phillipsia (Griffithides) portlockii meek and worthen, 1865, p. 268-270; 1873, p. 525-528, pi. 19, fig. 6.

Griffithides portlockii voodes, 1887, p. 93, pi. 3, fig. 9; walter, 1924, p. 321-322, pi. 27, figs. 17-19.

Phillipsia portlockii keyes, 1894, p. 236, pi. 32, fig. 7.

Exochops portlockii weller, 1936, p. 707, pi. 95, fig. 7; SHIMER AND SHROCK, 1944, p. 645, pi. 276, fig. 7; hahn and hahn, 1970, p. 194-195.

Griffithides salinensis brezinski, 1986, p. 141-142, figs. Ia-Ii.

Types.-Holotype from the Keokuk Limestone of Illinois (U of 111. X-104).

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Description.-see description in Meek and Worthen (1873) and Walter (1924).

Discussion.-Because the original illustration of this species is a line drawing, and because Weller (1936) failed to clearly discuss pygidial
characters that diagnose Exochops, there has been much confusion regarding the affinities of E. portlockii (Meek and Worthen) in particular and
the genus Exochops in general. Brezinski (1986) mistakenly assigned pygidia from the Keokuk Limestone of Missouri to a new species Griffithides
salinensis because of the previous shortcomings with illustrations of this species. That is why the holotype is reillustrated herein. Clearly,
Brezinski's (1986) specimens are conspecific with the type specimen of E. portlockii. More significantly, the pygidial characters that diagnose E.
portlockii are similar to the illustrated pygidium of Phillipsia meramecensis (Shumard, 1855, pi. B, fig. 9). The overall plan of the pygidium for
both species is similar, and they are known from the same stratigraphie interval, but the illustration of P. meramecensis is poor, and no type is
known. While it is highly likely that these two species are conspecific, there is no way to be certain.

EXOCHOPS BURLINGTONENSIS new species

Figure 3.14-3.20

Diagnosis.-Palpebral lobes highly elevated, small; rounded eyes small, reniform. Glabella rounded at base. Ll small, depressed.

Description.-Cephalon highly vaulted, dorsal furrow deep. Glabella strongly arched in transverse profile, moderately arched in longitudinal
profile, vertical to overhanging anterior margin, oval in outline, rounded at base, covered by small granular prosopon in frontal lobe, granules
becoming coarser to the posterior. Dorsal furrow deep. Ll small, depressed, subtriangular in outline. S1 shallow, straight, relatively broad.
Occipital ring wide, with small lateral lobes, descending into straight, wide occipital furrow. Anterior facial sutures straight, moderately
anteriorly diverging, sharply curved at â. Posterior facial suture with sharp curve at e and a short, straight, posteriorly diverging section.
Palpebral lobes highly elevated, short, crescentic in outline; palpebral suture semicircular between y and å. Eyes are small, narrow; eye socle
shallow, relatively broad. Lateral border broad, shallow, extending onto short, blunt genal spine; lateral margin rounded.

Pygidium outline semielliptical, vaulting moderate. Axis broad, low in transverse profile, nearly straight in longitudinal profile, composed of
more than 12 broad rings separated by narrow, deep ring furrows, bluntly rounded at terminus. Pleural fields arched in transverse profile,
flattened adaxially, steeply inclined and curved at margin, composed of 13 to IS narrow ribs that extend behind the axis. Pleural furrows narrow,
deep in the anterior.

Etymology.-Named for the Osagean Burlington Limestone from where it is known.

Types.-Holotype, a partial cephalon from the Burlington Formation, Columbia, Missouri, precise location unknown, UMC 17488. Paratype, an
incomplete pydigium from the Burlington Formation, center of section IS, T48N, R14W, CM 53904.

Occurrence.-Known from the Burlington Limestone (lower Osagean) of Boone, Pettis, and Cooper Counties, Missouri.

Discussion.-Exochops burlingtonensis n. sp. can be distinguished from E. portlockii by the former possessing a smaller, narrower palpebral lobe;
smaller eyes; a rounded base to the glabella; smaller, depressed Ll ; and broad and deep occipital furrow. The eyes of E. portlockii are large
semispherical mounds. While E. portlockii has a distinct subocular grove similar to E. burlingtonensis, the former species also has a row of
granules along the subocular ridge. The paratype pygidium of E. burlingtonensis has an elevated terminal ring similar to that of the type
species, but because of weathering it is difficult to ascertain whether the terminal ring protrudes like that of E. portlockii.

The highly vaulted cranidium and general plan of the pygidium of E. burlingtonensis is similar to Hesslerides bufo. However, the latter species
does not exhibit the elevated, narrow palpebral lobe, small eye, and depressed Ll that E. burlingtonensis possesses. Furthermore, the narrow,
poorly defined border of H. bufo allows distinction between the two taxa. Hesslerides arcentensis Brezinski 2000 and H. pustulosus differ from E.
burlingtonensis in possessing glabellae that exhibit bluntly rounded anterior and posterior termini, straight-sided glabellae between â and -y,
and narrower Sl.Exochops burlingtonensis also exhibits depressed Ll and considerably more glabellar vaulting than any of the species of
Hesslerides.

SPERGENASPIS Brezinski, 1987

SPERGENASPIS BCXJNENSIS new species

Figure 5.1-5.11

Diagnosis.-Anterior branch of facial sutures acute at â. Pygidial border posterior to axis broad, 0.25 the total pygidial length. Narrow pleural
fields comprised of 4 to 5 ribs.

Description.-Frontal glabellar lobe with very low relief and vaulting, subcylindrical, bluntly rounded in front, constricted at y. Anterior branch
of facial sutures moderately diverging, with a nearly straight, long section that is broadly rounded at â and meets á well away from the axial
line. Posterior facial suture short, diverging posteriorly. Sl shallow, faint. Ll oval. Anterior margin sharply recurved, upturned anterior to
glabelle with shallow border furrow extending laterally, intersecting sutures posterior of â. Palpebral lobes large, semicircular, flattened distally.
Dorsal furrow shallow, narrow, sinuous. Occipital furrow narrow, straight. Occipital ring flattened.

Pygidium outline parabolic, with little vaulting and relief, 0.7 times longer than wide. Axis low, evenly arched in tranverse profile, nearly

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straight in longitudinal profile, narrow, 0.3 the maximum pygidial width, comprising 0.75 of the total pygidial length, strongly posteriorly
tapering to a sharply rounded terminus, comprised of 9-10 narrow rings, separated by straight, shallow, narrow ring furrows. Pleural fields low,
evenly arched, composed of 4 to S ribs, each made up of anterior band slightly wider than the posterior band. Pleural furrows narrow, distinct;
interpleural furrows shallow, faint in anterior ribs, indistinct in posterior ribs. Border broad, 0.25 the pygidial width at axial line, smooth, low to
nearly flat, with faint impressions of the first pleural furrow.

Etymology.-Named for the Boone Formation of Oklahoma and Arkansas from where it is found.

Types.-Holotype, a nearly complete cranidium from the Boone Formation of Oklahoma, CM 53905. Paratypes, a partial cranidium and complete
pygidium from the same location as the holotype, CM 53906, 53907, and a complete testate pygidium from the same location, CM 53908. All
specimens from USGS Locality PC-1190.

Occurrence.-The Boone Formation of eastern Oklahoma and northern Arkansas, and the Burlington Formation of southwestern Missouri.

Discussion.-Spergenaspis boonensis n. sp. differs from its congeners by possessing a very wide pygidial border, more deeply incised pleural and
axial furrowing, fewer pygidial rings and ribs, and a more sharply posteriorly tapering axis. The partial cranidium of S. boonensis is very similar
to S. mauvaisensis in that the anterior branch of the facial sutures is nearly straight and only moderately anteriorly diverging, with a bluntly
rounded â. However, the angle at â is more acute in S. boonensis. Spergenaspis boonensis also appears to have glabellar constriction at y similar
to 5. mauvaisensis, but the incompleteness of the cranidium and exfoliation of the glabella make it difficult to make a definitive comparison.
Spergenaspis boonesis appears to be slightly older than either S. mauvaisensis or S. salemi; as best as can be determined, it is early to middle
Osagean.

Genus RICHTERELLA Hessler, 1965

RICHTERELLA CARTERI new species

Figure 5.12-5.14

Diagnosis.-Pygidium parabolic, low in relief and vaulting; axis narrow, low in relief and vaulting.

Description.-Pygidium low in relief and vaulting, outline parabolic. Axis short, narrow, 0.3 times the maximum pygidial width, mildly and
evenly arched in transverse profile, straight in longitudinal profile, mildly posteriorly tapering, sharply rounded at terminus, 0.8 the total
pygidial length, composed of 11 smooth, evenly wide rings. Ring furrows shallow, narrow, straight, shallower laterally, faint indications of
shoulders present. Pleural fields low, mildly arched in transverse and longitudinal profiles, composed of six to seven ribs. Ribs consist of broad,
slightly elevated anterior bands, and slightly narrower, lower posterior bands. Anterior pleural furrows extend onto border. Interpleural furrows
shallow on anterior two to three ribs, indistinct on posterior ribs. Rib definition becomes indistinct to the posterior. Border relatively wide, about
0.2 of the pygidial length at axial line. Margin rounded, underturned.

Etymology.-Named in honor of Dr. John L. Carter, collector of the holotype.

Types.-Holotype, a complete pygidium, CM 53909, from the Burlington Limestone, center of border between sections 4 and 9, T46N, R19W,
Cooper County, Missouri.

Occurrence.-A pygidium from the Burlington Formation, Sweeney Quarry, Cooper County, Missouri.

Discussion.-Richterella carteri n. sp. differs considerably from the type species R. snakedenensis. The latter is much more vaulted, especially in
the pleural fields, and the pleural furrows are distinct even on posterior ribs. Superficially, R. carteri appears more similar to Perexigupyge
chouteauensis than to the type species, because the latter species has low relief and vaulting and indistinct pleural ribs. However, R. carteri
differs from P. chouteauensis by having a narrow axis that is weakly posteriorly tapering, subdued axial and pleural furrowing, and a distinct
border. Consequently, while there remains a question as to whether R. carteri should be assigned to Richterella or Perexigupyge, currently
identified pygidial characters suggest that it is more appropriately assigned to Richterella.

ACKNOWLEDGMENTS

Lower Mississippian and especially Osagean trilobite ranges were greatly enhanced by specimens made available for study by J. H. Stitt, J. F.
Miller, J. D. Loch, R. A. Garney, J. L. Carter, A. D. Kollar, J. F. Taylor, C. A. Kertis, and J. Golden. The manuscript was improved by reviewers B.
S. lieberman and R. M. Owens. Their suggestions are very much appreciated.

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ACCEPTED 25 JANUARY 2006

DAVID K. BREZINSKI

Maryland Geological Survey, 2300 St. Paul Street, Baltimore, 21218

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