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Journal of Paleontology: NEW TRILOBITE TAXA FROM UPPER...http://findarticles.com/p/articles/mi_qa3790/is_200405/ai_n937...1 of 98/12/2008 12:21 AM
FindArticles>Journal of Paleontology >May 2004> Article> Print friendly 
Loch, James D ABSTRACT-Previously undescribed trilobites associated with microbial patch reefs have been recovered from the Upper Cambrian (upperSteptoean Stage) Ore Hill Limestone Member of the Gatesburg Formation in south-central Pennsylvania. Reefs in the Imler Quarry andDrab-Beaverton measured sections yielded low-diversity assemblages of trilobites that we assign to the uppermost Cliffia lataegenae Subzone of the Elvinia Zone. The faunas are dominated by the catillicephalid Buttsia drabensis Wilson, 1951, and include several new taxa that are notknown from coeval off-reef facies. Fine-scale correlation between the two sections reveals that the reefs in the Drab-Beavertown section areslightly younger than those at Imler Quarry and represent the youngest Steptoean fauna described from the Appalachian region.New taxa include Imlerella n. gen. (type species Imlerella praecipita n. sp.), Stittaspis n. gen. (type species Stittaspis loria n. sp.), and Dellearogersi n. sp. Stigmacephalusl distorta Wilson, 1951, is reassigned to Stittaspis and is restricted to the type. Additional material of Buttsiadrabensis illustrates features not discernible in previous treatments of this variable species.INTRODUCTIONIN AN earlier paper (Taylor et al., 1999), we reported the serendipitous discovery of an atypical assemblage of Elvinia Zone (upper Steptoean)trilobites from microbial reefs in the Ore Hill Member of the Gatesburg Formation from south-central Pennsylvania. The purpose of that paper was to compare the previously undescribed reef biofacies (sensu Ludvigsen and Westrop, 1983) with coeval nonreef faunas, primarily at thegenus level, and describe one of several new species in the reef fauna, Cheilocephalus quadratus Taylor and Loch in Taylor, et al., 1999. Here wedescribe two new genera and three new species from the same and from a slightly younger reef interval discovered more recently. The status of the genus Eshelmania Wilson, 1951, and the range of variability in Buttsia drabensis Wilson, 1951, are also discussed.BIOSTRATIGRAPHIC CONTEXTThe first reef-related trilobite assemblage recognized in the Ore Hill Member (Taylor, et al., 1999) was recovered from the Imler Quarry section,near the south end of the unit's outcrop belt (Fig. 1, see Wilson, 1951, 1952 for additional locality information). We assigned the faunas recoveredfrom the microbial boundstones and inter-reef grainstones at that locality (Fig. 2; IQ-25.5, IQ-26, IQ-27) to the Cliffia lataegenae Subzone of theElvinia Zone based upon the occurrence (albeit rare) of Deckera completa Wilson, 1951, and Cliffia lataegenae (Wilson, 1949). We havepreviously demonstrated that these two species are restricted to that subzone (Loch and Taylor, 1995). The reef-related collections from ImlerQuarry are dominated by the catillicephalid Buttsia drabensis, Pseudosaratogia spp., and a previously undescribed species of Dellea Wilson, 1949(D. rogersi n. sp., described herein) (Table 1). The collections also include a species whose unusual, strongly vaulted cephalon requires erection of a new genus and species, Imlerella praecipita. More recent fieldwork has revealed that trilobite-bearing microbial reefs also occur near the top of the Cliffia lataegenae Subzone in the Drab-Beavertown section, approximately 50 km north of Imler Quarry (Fig. 1). Sizeable collections fromthese newly discovered reefs expand the known geographic and stratigraphie ranges of Cheilocephalus quadratus, Dellea rogersi n. sp., andlmlerella praecipita n. gen. and sp. Like those from Imler Quarry, collections from reefs high in the Cliffia lataegenae Subzone in theDrab-Beavertown section (collections DB-95 and DB-113; Fig. 1) lack many of the common species of the Elvinia Zone and are, rather, dominated by Buttsia drabensis (Table 1). The Drab-Beavertown reefs, however, include another previously undescribed taxon, Stittaspis loria n. gen. andsp., that does not occur in the reefs to the south.The relative ages of the Imler Quarry and Drab-Beaverton microbial reefs have been difficult to establish, in part because the former areimmediately overlain by a unit of nonfossiliferous dolomite that occurs in the middle of the Ore Hill Member in the southernmost Pennsylvaniasections (Wilson, 1951, 1952). Consequently, the position of the Steptoean-Sunwaptan boundary (base of the Irvingella major Subzone) withinthese sections remains poorly constrained. In contrast, we were able to establish the position of that stadial boundary with much greater precisionin the Drab-Beavertown section, demonstrating that the reefs that contain Stittaspis loria lie within the highest 7-8 m of the Cliffia lataegenaeSubzone (Fig. 2). These reefs also yielded Cheilocephalus quadratus and Imlerella praecipita n. gen. and sp., which occur in the reefs at ImlerQuarry, but lack Dellea rogersi n. sp., despite recovery of over 150 specimens. Dellea rogersi n. sp. was recovered, however, from poorly exposedthrombolitic boundstone almost 3 m lower in the section, suggesting that this lower reefal horizon (DB-86) is equivalent in age to the reefs atImler Quarry (Fig. 2). Additional support for the correlation of the reef(s) at DB-86 with those at Imler Quarry is provided by the occurrence in both of Drabia acroccipita Wilson, 1951, a species restricted to a fairly narrow stratigraphie interval in the middle of the Cliffia lataegenaeSubzone (Loch and Taylor, 1995). We conclude, therefore, that the reefs containing Stittaspis loria n. gen. and sp. are slightly younger than thoseat Imler Quarry, making this new species useful for assignment of strata that yield it to the very highest part of the Sunwaptan Stage, above therange of Drabia acroccipita and Dellea rogersi n. sp. (Fig. 2).SYSTEMATIC PALEONTOLOGY Morphologic terms used are those recommended by Whittington (1997). All types and illustrated specimens are reposited at the CarnegieMuseum of Natural History (CM) in Pittsburgh, Pennsylvania. Occurrence data are keyed by measured section abbreviation and measureddistance (in feet) above the base of the section. Abundance data are recorded as (cranidia-pygidialibrigenae) for each collection horizon.
Journal of Paleontology: NEW TRILOBITE TAXA FROM UPPER...http://findarticles.com/p/articles/mi_qa3790/is_200405/ai_n937...2 of 98/12/2008 12:21 AM
Superfamily DIKELOCEPHALACEA Miller, 1889Family PTYCHASPIDIDAE Raymond, 1924Genus IMLERELLA new genusType species.-Imlerella praecipita new species, by monotypy and original designation, herein, from Ore Hill Member of the Gatesburg Formation,south-central Pennsylvania.Diagnosis.-A genus of Ptychaspididae with a semicircular cranidium of small to moderate size, dominated by subrectangular glabella with afaintly impressed transglabellar is furrow; moderately convex (tr.), strongly declined to overhung preglabellar area; narrow, laterally taperinganterior border poorly defined by very faint border furrow and concentration of coarse punctae on border; faint occipital furrow; broad anteriorfields and palpebral areas defined laterally by subparallel anterior facial sutures, posterolateral projection recurved posteriorly to produce shortposteriorly directed spine; straight posterior border furrow. Librigenae subtriangular in outline, narrow (tr.), with massive, bluntly pointedgenal spine covered with coarse terrace ridges; broad, nearly vertical lateral border defined by faint to moderately impressed border furrow, widened slightly posteriorly, with coarse terrace ridges on all but most proximal posterior portion. Nearly vertical librigenal doublure extendsinward (upward) approximately to lateral border furrow.Etymology.-Named for the Imler Quarry from which the first specimens were recovered.Discussion.-Imlerella n. gen. is tentatively assigned to the Ptychaspididae based upon the presence of a subrectangular glabellar outline, atransglabellar Is glabellar furrow, and small palpebral lobes that stand in low relief above the palpebral areas. Westrop (1986) used thesecharacters in his diagnosis and discussion of the Ptychaspididae. Further, we note the presence of coarse terrace ridges on the librigenareminiscent of the prosopon on many ptychaspidid species.The subquadrate cranidium, rectangular glabella, wide fixigenae, anteriorly centered palpebral lobes, overturned preglabellar area, andmorphology of the posterior border of the cranidium are nearly unique among Upper Cambrian trilobites. Tholifrons Palmer, 1968, appearssimilar in glabellar shape, fixigenal proportions, shape of the librigena, and position of the palpebral lobes. The preglabellar area of Tholifrons,however, is gently to moderately declined, rather than overhung. Further, the short posterior border spine of Imlerella n. gen. contrasts with asimple triangular border on Tholifrons. Paraphoreotropis Shaw (in Clark and Shaw, 1968) also shares the rectangular glabella, broad fixigenae,overturned preglabellar area, and anteriorly positioned palpebral lobes of Imlerella n. gen. The cranidium of Paraphoreotropis, as noted by Pratt(1992), has a well-defined anterior border furrow that defines a horizontal anterior border. The geometry of the border separatesParaphoreotropis from both Tholifrons and Imlerella n. gen. There is sufficient similarity between these three genera to suggest that they may represent a subfamily within the Ptychaspididae. We decline to erect a subfamily at this time, however, because of the absence of a pygidium foreither Imlerella n. gen. or Paraphoreotropis.IMLERELLA PRAECIPITA new speciesFigure 4.1-4.14Genus and species undetermined in TAYLOR, LOCH, AND PERFETTA, 1999, fig. 5.8-5.10.Diagnosis.-As for the genus.Description.-Cranidium moderate in size, up to 6.0 mm in sagittal length, semicircular in outline, strongly convex (sag.); glabella long,subrectangular, broadly rounded anteriorly, moderately convex (tr.), weakly keeled in larger individuals, steeply declined anteriorly approaching preglabellar furrow; axial furrows well impressed, broad except between where very deeply impressed between Is furrow and faintocular ridge; preglabellar furrow narrow and more weakly impressed. Two sets of lateral glabellar furrows; 1s-posteriorly directed, faintly impressed laterally, extending transversely across axis as very faintly impressed transglabellar furrow in some small specimens, 2s-orientedtransversely, located opposite posterior end of palpebral lobes, very faintly impressed to obsolete on small specimens, faintly impressed on largerspecimens. Occipital furrow broad, faintly to moderately impressed, straight over axis, bifurcating near axial furrows; occipital ring short (sag.),subrectangular to subovoid in outline, tapering slightly adaxially, moderately convex (tr.), elevated above posterior end of glabella. Preglabellararea moderately convex (sag.), declined to appear vertical to slightly overturned at anterior margin, very faint anterior border furrow on somespecimens defines short, laterally tapering, coarsely punctate anterior border; preglabellar field long, approximately two to three times length of  border when viewed anteriorly. Preocular field broad (tr.), moderately convex, steeply declined; palpebral areas moderately wide, slightly inflated, delimited anteriorly by faint eye ridge; posterior area long (exsag.), triangular, moderately convex, subhorizontal. Posterior bordernarrow (exsag.), of uniform width along proximal two-thirds, expanded slightly and directed posteriorly as short tubular spine along distalone-third; posterior border furrows faintly to moderately well impressed, gently curved posteriorly, intersect axial furrows. Palpebral lobessimple, flaplike, with very faint palpebral furrows, centered anterior to glabellar midlength. Anterior branch of facial sutures subparallel toslightly divergent in front of palpebral lobe, curved adaxially approaching anterior margin; posterior branch of facial suture moderately divergent, continued in broad curve to intersect posterior margin at obtuse angle. Prosopon smooth.Librigena elongate, triangular in outline; smooth and moderately wide (tr.) genal field of uniform width, gently convex, and steeply declinedaway from low eye socle; eye socle furrow distinct but weakly impressed. Very gently convex, nearly vertical lateral border approximately 
Journal of Paleontology: NEW TRILOBITE TAXA FROM UPPER...http://findarticles.com/p/articles/mi_qa3790/is_200405/ai_n937...3 of 98/12/2008 12:21 AM
one-half width of genal field, defined by moderately impressed border furrow that fades posteriorly before intersecting posterior border furrow.Posterior border furrow short, slightly to moderately impressed, directed posteriorly abaxially; posterior border poorly defined, merges abaxially  with genal spine. Genal spine massive, over two-thirds length (exsag.) of genal field, narrow (tr.), bluntly pointed, covered with coarse terraceridges.Pygidium unknown.Etymology.-From praecipitium (Latin), cliff or precipice, in reference to the steeply declined preglabellar area.Types.-Holotype cranidium CM 52024. Paratype cranidia CM 52025-27, 52029; paratype librigenae CM 52028, 52030-31. Additional materialincluded 13 cranidia and three librigenae.Occurrence.-Most common in microbial reef boundstones and related grainstones located high in the Cliffia lataegenae Subzone of the ElviniaZone (uppermost Steptoean Stage). Imler Quarry section: IQ-25.5 (2-0-1), IQ-26 (6-0-3). Drab-Beaverton section: DB-59 (2-0-0), DB-95 (7-0-2).Discussion.-Association of the cranidium with librigenae for this species was based upon complimentary geometry of the facial sutures and the very faint impression of border furrows anteriorly on the cranidium and laterally on the librigena.Superfamily OLENACEA Burmeister, 1843Family ELVINIIDAE Kobayashi, 1935Subfamily DOKIMOCEPHALINAE Kobayashi, 1935Genus DELLEA Wilson, 1949Type species.-Dellea wilbernsensis Wilson, 1949, by original designation, from the Morgan Creek Member of the Wilberns Formation of centralTexas (see Ludvigsen and Westrop, 1983, p. 16-17 for discussion).Discussion.-The diagnosis provided for Dellea by Ludvigsen and Westrop (1983) is followed here with two emendations. We note the presence of alaterally tapering anterior border for Dellea. Further, addition of D. rogersi n. sp. expands the generic concept to include at least one species withshallow, rather than deeply impressed, axial and preglabellar furrows and faint to indistinct palpebral furrows.The number and identity of Dellea species represented in the Ore Hill fauna has been debated and remains unresolved. Wilson (1951) reportedthree species of Dellea from the Ore Hill Member: D. butlerensis, Frederickson (1949), D. saratogensis (Resser, 1942), and D. suada (Walcott,1890). Among these he cited D. suada as the most common trilobite in the Elvinia Zone in the Ore Hill. However, he applied a very broadmorphologic concept for D. suada. Thus he synonymized D. wilbernsensis (the type species from the Wilberns Formation in Texas [Wilson, 1949]) with D. suada. A similarly broad concept was used for D. suada in studies of Elvinia Zone faunas from Montana (Grant, 1965), Oklahoma (Stitt,1971), and Missouri (Kurtz, 1975). In each of these studies the material assigned to Dellea was interpreted as representing one long-ranging andrather highly variable species. In contrast, Palmer (1965) argued that D. wilbernsensis and D. suada can be distinguished as separate species with granular versus smooth prosopon, respectively. Ludvigsen and Westrop (1983), in redescribing type material for D. saratogensis from theGalway Formation in New York, echoed Palmer's concern that prosopon had received insufficient attention in earlier diagnoses. They questionednot only Wilson's synonymy of D. wilbernsensis with D. suada, but also his synonymy of D. glabellamersa Wilson, 1949, (another species fromthe Wilberns Formation) with D. saratogensis. They noted that the specimens that Wilson (1951) had illustrated from the Ore Hill as D.saratogensis lack the coarse granular prosopon displayed by all specimens from the Galway Formation. They concluded that the Ore Hillmaterial represents a different species, perhaps D. glabellamersa. Granular cranidia that can confidently be assigned to D. saratogensis werelater reported from the Ore Hill (Taylor et al., 1999), but only from the Pseudosaratogia magna Subzone at the base of the member. A moredefinitive appraisal of the status of D. wilbernsensis and D. glabellamersa will require thorough reevaluation of these species. The distribution of these granulose morphs, along with documentation of any other morphologic features, in the Wilberns Formation and the Ore Hill Limestone will be necessary to justify their retention as distinct species. Finally, Palmer (1965) assigned D. butlerensis to the genus Apachia Frederickson, 1949. We note that the convexity of the glabella (tr., sag.) is not as pronounced as in the type species of Apachia (A. trigonis Frederickson, 1949), theelevation of the glabella above the occipital ring is not as great, and the glabella is not as strongly tapered. We do not agree with Palmer's (1965)action and retain D. butlerensis within the genus. Another unresolved issue that bears on the question of how many Dellea species occur in the Ore Hill fauna is the status of the genus Eshelmania Wilson. Wilson (1951) erected this genus to accommodate two uncommon species from the Ore Hill Member and equivalent strata in theConococheague Formation. Despite its similarity to Dellea, Wilson considered Eshelmania a separate genus characterized by a longer, morequadrate glabella, well-defined lateral furrows, and strongly recurved posterior limbs. Grant (1965) synonomized Eshelmania with Dellea,arguing that the glabellar proportions, curvature of the posterior margin of the fixigenae, and degree of impression of the glabellar furrows wereinsufficient as generic characters to separate the two concepts. However, Grant's arguments focused on each trait in isolation and did not considertheir combined potential for denning the genus. We disagree with Grant (1965) and subsequent authors who accepted his synonymy (e.g., Westrop, 1986) and retain Eshelmania as a distinct genus, adding the form of the anterior border to the diagnosis. The type species, Eshelmaniasnoburgensis Wilson (1951, p. 642, pi. 92, figs. 9-15), shows pronounced rearward extension of the posterolateral projection (resulting fromstrongly posteriorly curved posterior border furrows and distally expanded posterior borders) and an anterior border that does not taper

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